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Morphological Analysis of Tropical Bulbs and Environmental Effects on Flowering and Bulb Development of Habranthus and Z...

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PAGE 1

MORPHOLOGICAL ANALYSIS OF TROPICAL BULBS AND ENVIRONMENTAL EFFECTS ON FLOWER ING AND BULB DEVELOPMENT OF Habranthus robustus AND Zephyranthes spp By CAMILA BRITO PAULA A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLOR IDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2006

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Copyright 2006 by Camila Brito Paula

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This document is dedicated to my hus band and to my parents in Brazil.

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iv ACKNOWLEDGMENTS I thank Dr. Rick Schoellhorn for giving me the opportunity of entering the “plant world,” for believing that an architect could be able to learn some science and for serving as my major advisor for the first half of th e process of my research. I thank Dr. Dennis McConnell for playing a large role in my research from the beginning, for making my “bulb-killing” experience much more enjoya ble, for accepting the challenge of becoming the chair of my committee in the middle of the process, and also for his enormous patience and willingness to help me during th e preparation of this thesis. I thank Dr. Wagner Vendrame for helping me start this w hole process and for much advice. I also thank Dr. Alan Meerow for serving on my supervisory committee. I thank Mrs. Fe Almira for her immense he lp with the images of this thesis, Mrs Carolyn Bartuska for her help with experi ments set ups and statistical analysis, and Robert Weidman and his crew fo r all their greenhouse assistance. I would like to thank my husband for the encouragement, the emotional and “technical” support, for his love and friends hip. And I thank my mom and dad for their unconditional love, “long-distance” support and for giving me the educational and emotional base that allowed me to achieve my dreams. Lastly, I would like to thank all the friends I made in Gainesville, especially Carmen Valero-Aracama.

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v TABLE OF CONTENTS page ACKNOWLEDGMENTS.................................................................................................iv LIST OF TABLES............................................................................................................vii LIST OF FIGURES.........................................................................................................viii ABSTRACT....................................................................................................................... xv CHAPTER 1 INTRODUCTION........................................................................................................1 2 LITERATURE REVIEW.............................................................................................6 Aerial Organs and Tissues............................................................................................8 Underground Tissues....................................................................................................8 Taxonomy and Origin of Geophytes............................................................................9 Geophytes Growth and Development...........................................................................9 Dormancy...................................................................................................................10 Flowering Process.......................................................................................................12 Anatomy and Physiology of Flower Initiation....................................................14 Factors Affecting Flower Initiation.....................................................................17 Photoperiod..................................................................................................18 Light quality and quantity............................................................................18 Temperature.................................................................................................19 Vernalization................................................................................................20 Flower Initiation Process in Bulbous Plants........................................................21 Flower Bulb Cultivation.............................................................................................21 Flower Bulb Forcing...................................................................................................22 Tropical Bulbs and Amaryllidaceae...........................................................................24 Hippeastrum spp.........................................................................................................25 Scadoxus multiflorus – Blood Lily.............................................................................26 Agapanthus africanus – African Lily.........................................................................27 Habranthus robustus and Zephyranthes spp. – Rain Lilies........................................28 3 BULB MORPHOLOGY............................................................................................31 Comparative Study.....................................................................................................31

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vi Materials and Methods........................................................................................31 Results and Discussion........................................................................................33 Bulb type and size........................................................................................33 Leaf arrangement and morphology..............................................................37 Floral initiation.............................................................................................41 Meristematic region......................................................................................48 General anatomical and morphological arrangement...................................55 Experiment 1...............................................................................................................60 Materials and Methods........................................................................................61 Results.................................................................................................................61 4 ENVIRONMENTAL EFFECTS – FERTIGATION FREQUENCY AND FERTILIZER RATES ON FLOWERING IN Habranthus robustus AND Zephyranthes spp .......................................................................................................64 Experiment 1...............................................................................................................67 Materials and Methods........................................................................................67 Results and Discussion........................................................................................69 Experiment 2...............................................................................................................84 Materials and Methods........................................................................................84 Results and Discussion........................................................................................86 5 ENVIRONMENTAL EFFECTS – LIGHT LEVELS ON FLOWERING AND BULB DEVELOPMENT IN Habranthus robustus AND Zephyranthes SPP...........93 Materials and Methods...............................................................................................94 Experiment 1.......................................................................................................94 Experiment 2.......................................................................................................96 Results and Discussion...............................................................................................97 Experiment 1.......................................................................................................97 Experiment 2.....................................................................................................100 6 CONCLUSIONS......................................................................................................113 LIST OF REFERENCES.................................................................................................119 BIOGRAPHICAL SKETCH...........................................................................................127

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vii LIST OF TABLES Table page 3-1 Tropical geophytes used in the morphological studies............................................32 3-2 Geophyte size and characteri stics of plants examined.............................................34 3-3 Leaf size of plants examined....................................................................................39 3-4 Relationship between leaf and flower formation, and number of flowers per inflorescence of plants examined.............................................................................43 4-1 Mean bulb size and weight prior to experiment.......................................................85 5-1 Photometric readings and temperatur es at three locations / treatments...................96 5-2 Rainfall monthly summary in inches for Gainesville area in 2004 and 2005 according to the Florida Automated Weather Network...........................................97

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viii LIST OF FIGURES Figure page 3-1 From left to right Zephyranthes, Habranthus, Hippeastrum Scadoxus bulbs, and immature Agapanthus rhizome................................................................................35 3-2 Sections of Zephyranthes bulbs................................................................................35 3-3 Sections of Habranthus bulbs..................................................................................36 3-4 Sections of Hippeastrum bulbs................................................................................36 3-5 Sections of Scadoxus bulbs......................................................................................36 3-6 Longitudinal sections of an Agapanthus rhizome....................................................37 3-7 Hippeastrum leaf arrangement.................................................................................39 3-8 Habranthus leaf arrangement...................................................................................39 3-9 Zephyranthes leaf arrangement................................................................................40 3-10 Agapanthus leaf arrangement...................................................................................40 3-11 Scadoxus leaf arrangement.......................................................................................40 3-12 Longitudinal sections of a Hippeastrum bulb..........................................................43 3-13 Sections of Habrantus bulbs....................................................................................43 3-14 Sections of Zephyranthes bulbs................................................................................44 3-15 Cross section of a Habranthus bulb.........................................................................44 3-16 Longitudinal section of a Habranthus bulb..............................................................45 3-17 Habranthus bulb with three flower stalks................................................................45 3-18 Development of a Habranthus flower......................................................................46 3-19 Sections of Scadoxus bulbs......................................................................................46

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ix 3-20 Flower stalk emerged a nd pseudo-stem arising from a Scadoxus bulb....................47 3-21 Sections of Agapanthus rhizomes............................................................................47 3-22 Photomicrograph of the meristematic region of a Hippeastrum bulb......................50 3-23 Photomicrograph of the meristematic region of a Hippeastrum bulb......................50 3-24 Photomicrograph of the meristematic region of a Habranthus bulb........................51 3-25 Photomicrograph of the meristematic region of a Habranthus bulb........................51 3-26 Photomicrograph of the meristematic region of a Zephyranthes bulb.....................52 3-27 Photomicrograph of the meristematic region of a Zephyranthes bulb.....................52 3-28 Photomicrograph of the meristematic region of a Scadoxus bulb............................53 3-29 Photomicrograph of the meristematic region of a Scadoxus bulb............................53 3-30 Photomicrograph of the meristematic region of an Agapanthus rhizome................54 3-31 Photomicrograph of the meristematic region of an Agapanthus rhizome................54 3-32 Longitudinal section of a Hippeastrum bulb showing its general anatomy and morphology..............................................................................................................55 3-33 Longitudinal section of a Hippeastrum bulb showing its general anatomy and morphology..............................................................................................................55 3-34 Longitudinal section of a Zephyranthes bulb showing its general anatomy and morphology..............................................................................................................56 3-35 Longitudinal section of a Zephyranthes bulb...........................................................56 3-36 Sections of Habranthus bulbs..................................................................................57 3-37 Longitudinal section of a Scadoxus bulb showing its general anatomy and morphology..............................................................................................................57 3-38 Longitudinal section of a Scadoxus bulb showing its general anatomy and morphology..............................................................................................................58 3-39 Longitudinal section of an Agapanthus rhizome showing its general anatomy and morphology........................................................................................................59 3-40 Longitudinal section of an Agapanthus rhizome showing its general anatomy and morphology........................................................................................................59

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x 3-41 Number of leaves and flowers produced by Habranthus bulbs in 2005..................63 3-42 Number of leaves and flowers produced by Zephyranthes bulbs in 2005...............63 4-1 Habranthus bulbs during fertigation experiment with different plastic tags in different color distinguishing the three treatments...................................................68 4-2 Habranthus bulbs being weighed after completion of experiment..........................69 4-3 Number of flowers of Habranthus robustus bulbs in year 2 as affected by fertigation frequency................................................................................................74 4-4 Number of leaves of Habranthus robustus bulbs in year 1 as affected by fertigation frequency................................................................................................74 4-5 Number of offsets of Habranthus robustus bulbs in year 1 as affected by fertigation frequency................................................................................................75 4-6 Bulb size of Habranthus robustus in year 1 as affected by fertigation frequency...75 4-7 Total weight of Habranthus robustus bulbs in year 1 as affected by fertigation frequency..................................................................................................................75 4-8 Bulb weight of Habranthus robustus in year 1 as affected by fertigation frequency..................................................................................................................76 4-9 Number of flower buds of Habranthus robustus bulbs in year 1 as affected by fertigation frequency................................................................................................76 4-10 Number of leaves of Habranthus robustus bulbs in year 2 as affected by fertigation frequency................................................................................................76 4-11 Number of offsets of Habranthus robustus bulbs in year 2 as affected by fertigation frequency................................................................................................77 4-12 Bulb size of Habranthus robustus in year 2 as affected by fertigation frequency...77 4-13 Total bulb weight of Habranthus robustus in year 2 as affected by fertigation frequency..................................................................................................................77 4-14 Bulb weight of Habranthus robustus in year 2 as affected by fertigation frequency..................................................................................................................78 4-15 Number of flower buds of Habranthus robustus bulbs in year 2 as affected by fertigation frequency................................................................................................78 4-16 Total number of flowers on Habranthus and Zephyranthes bulbs from July to December 2004........................................................................................................79

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xi 4-17 Number of flowers of Zephyranthes spp bulbs in year 1 as affected by fertigation frequency................................................................................................79 4-18 Number of flowers of Zephyranthes spp bulbs in year 2 as affected by fertigation frequency................................................................................................80 4-19 Number of leaves of Zephyranthes spp. bulbs in year 1 as affected by fertigation frequency..................................................................................................................80 4-20 Number of offsets of Zephyranthes spp. bulbs in year 1 as affected by fertigation frequency..................................................................................................................81 4-21 Bulb size of Zephyranthes spp. in year 1 as affected by fertigation frequency.......81 4-22 Total fresh weight of Zephyranthes spp. bulbs in year 1 as affected by fertigation frequency..................................................................................................................81 4-23 Bulb weight of Zephyranthes spp. in year 1 as affected by fertigation frequency...82 4-24 Number of flower buds of Zephyranthes spp. bulbs in year 1 as affected by fertigation frequency................................................................................................82 4-25 Number of leaves of Zephyranthes spp. bulbs in year 2 as affected by fertigation frequency..................................................................................................................82 4-26 Number of offsets of Zephyranthes spp. bulbs in year 2 as affected by fertigation frequency..................................................................................................................83 4-27 Bulb size of Zephyranthes spp. in year 2 as affected by fertigation frequency.......83 4-28 Total fresh weight of Zephyranthes spp. bulbs in year 2 as affected by fertigation frequency..................................................................................................................83 4-29 Bulb weight of Zephyranthes spp. in year 2 as affected by fertigation frequency...84 4-30 Number of flower buds of Zephyranthes spp. bulbs in year 2 as affected by fertigation frequency................................................................................................84 4-31 Data points, regression lines, equations and coefficient of determination of number of leaves of Habranthus ..............................................................................88 4-32 Data points, regression lines, equations and coefficient of determination of number of offsets of Habranthus .............................................................................88 4-33 Data points, regression lin es, equations and coefficient of determination of bulb size of Habranthus ...................................................................................................89 4-34 Data points, regression lin es, equations and coefficient of determination of total fresh bulb weight of Habranthus .............................................................................89

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xii 4-35 Data points, regression lin es, equations and coefficient of determination of fresh bulb weight of Habranthus ......................................................................................89 4-36 Data points, regression lines, equations and coefficient of determination of number of flower buds of Habranthus .....................................................................90 4-37 Data points, regression lines, equations and coefficient of determination of number of leaves of Zephyranthes ...........................................................................90 4-38 Data points, regression lines, equations and coefficient of determination of number of offsets of Zephyranthes ...........................................................................90 4-39 Data points, regression lin es, equations and coefficient of determination of bulb size of Zephyranthes .................................................................................................91 4-40 Data points, regression lin es, equations and coefficient of determination of total fresh bulb weight of Zephyranthes ...........................................................................91 4-41 Data points, regression lin es, equations and coefficient of determination of fresh bulb weight of Zephyranthes ....................................................................................91 4-42 Data points, regression lin es, equations and coefficient of determination of fresh bulb weight of Zephyranthes ....................................................................................92 5-1 Plants under three different treatments full sun.......................................................96 5-2 Number of flowers of Habranthus robustus bulbs in year 2 as affected by light levels.......................................................................................................................10 2 5-3 Number of leaves of Habranthus robustus bulbs in year 1 as affected by light levels.......................................................................................................................10 3 5-4 Number of offsets of Habranthus robustus bulbs in year 1 as affected by light levels.......................................................................................................................10 3 5-5 Bulb size of Habranthus robustus in year 1 as affect ed by light levels.................103 5-6 Total weight of leaves of Habranthus robustus bulbs in year 1 as affected by light levels..............................................................................................................104 5-7 Bulb weight of Habranthus robustus in year 1 as affected by light levels............104 5-8 Number of flower buds of Habranthus robustus bulbs in year 1 as affected by light levels..............................................................................................................104 5-9 Root condition of Habranthus bulbs, under different treatments, after completion of experiment 1....................................................................................105

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xiii 5-10 Number of leaves of Habranthus robustus bulbs in year 2 as affected by light levels.......................................................................................................................10 5 5-11 Number of offsets of Habranthus robustus bulbs in year 2 as affected by light levels.......................................................................................................................10 5 5-12 Bulb size of Habranthus robustus in year 2 as affect ed by light levels.................106 5-13 Total weight of Habranthus robustus bulbs in year 2 as affected by light levels..106 5-14 Bulb weight of Habranthus robustus in year 2 as affected by light levels............106 5-15 Number of flower buds of Habranthus robustus bulbs in year 2 as affected by light levels..............................................................................................................107 5-16 Number of flowers of Zephyranthes spp bulbs in year 1 as affected by light levels.......................................................................................................................10 7 5-17 Number of flowers of Zephyranthes spp bulbs in year 2 as affected by light levels.......................................................................................................................10 8 5-18 Number of leaves of Zephyranthes spp bulbs in year 1 as affected by light levels.......................................................................................................................10 8 5-19 Number of offsets of Zephyranthes spp bulbs in year 1 as affected by light levels.......................................................................................................................10 8 5-20 Bulb size of Zephyranthes spp in year 1 as affected by light levels.....................109 5-21 Total weight of Zephyranthes spp bulbs in year 1 as a ffected by light levels......109 5-22 Bulb weight of Zephyranthes spp in year 1 as affect ed by light levels.................109 5-23 Number of flower buds of Zephyranthes spp bulbs in year 1 as affected by light levels.......................................................................................................................11 0 5-24 Number of leaves of Zephyranthes spp bulbs in year 2 as affected by light levels.......................................................................................................................11 0 5-25 Number of offsets of Zephyranthes spp bulbs in year 2 as affected by light levels.......................................................................................................................11 0 5-26 Bulb size of Zephyranthes spp in year 2 as affected by light levels.....................111 5-27 Total weight of Zephyranthes spp bulbs in year 2 as a ffected by light levels......111 5-28 Bulb weight of Zephyranthes spp in year 2 as affect ed by light levels.................111

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xiv 5-29 Number of flower buds of Zephyranthes spp bulbs in year 2 as affected by light levels.......................................................................................................................11 2

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xv Abstract of Thesis Presen ted to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science MORPHOLOGICAL ANALYSIS OF TROPICAL BULBS AND ENVIRONMENTAL EFFECTS ON FLOWER ING AND BULB DEVELOPMENT OF Habranthus robustus AND Zephyranthes spp By Camila Brito Paula May 2006 Chair: Dennis B. McConnell Major Department: Horticultural Sciences Morphologies of Hippeastrum x hybridum Habranthus robustus Zephyranthes spp, Scadoxus multiflorus and Agapanthus africanus were compared and contrasted. The first three species were true tunicate bulbs w ith similar shapes and distinct sizes, leaves emerged alternately from the center of the bul b, flower buds were formed alternately at the apical meristem (in a line in bulb cross se ction), and there were four leaves between each flower formed. Scadoxus was a true tunicate bulb with a thick rhizomatous structure at the base, thick scales, leaves arising in a pseudostem and flower buds formed centrally at the apical meristem. Its size is equivalent to Hippeastrum bulbs. Agapanthus is a rhizome with leaves arising alternately from lateral meristems, and flower buds formed centrally at the apical meristem. Habranthus and Zephyranthes had distinct patterns of leaf production. Habranthus bulbs had fewer leaves emerging during flowering compared to Zephyranthes which had a greater number of leaves throughout the entire year.

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xvi The responses of Habranthus robustus and Zephyranthes spp. to fertigation frequencies and fertilizer rates were ex amined. Experiment 1 tested fertigation frequencies of twice a week, once a week and every other week in both species. Regimens of once or twice a week were effective for flowering of both species All bulb development factors investigated incr eased as fertigation increased in Habranthus and Zephyranthes except for number of offsets on both species and number of leaves in Zephyranthes Experiment 2 tested bulb development res ponses when both species were fertilized with 20-10-20 at rates of 0, 75, 150, 300 ppm N. Habranthus bulbs treated with 75 and 150 ppm N had the greatest number of leaves and flower buds, and were larger and heavier, but bulbs treated with 300 ppm had more offsets. Zephyranthes bulbs treated with 150 ppm N had the greatest number of l eaves, offsets and flower buds; bulbs treated with 300 ppm N were largest. Bulb weight was similar in bulbs treated with 75, 150 or 300 ppm N. The responses of Habranthus robustus and Zephyranthes spp. to light levels of full sun, 30% and 60% shade were examined. Results demonstrated that both Habranthus and Zephyranthes flowered for a longer period under fu ll sun and 30% shade than 60% shade but plants had more flowers under 30% shade. All bulb development factors investigated increased as light level increased in both species. From this study it was concluded that pr eferred conditions for both species were fertigation of twice a week using fe rtilizer rate of 150 ppm N under full sun.

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1 CHAPTER 1 INTRODUCTION Bulbous plants are plants with a self-c ontained and highly developed food-storage mechanism that allows them to live unde rground. These plants are found all around the world; however the principal areas that are natural common habitats of a high percentage of bulbous plants are between 23o North and 45oSouth latitudes (Du Plessis and Duncan, 1989) in the Mediterranean, South Africa, Middl e East and the Pacific seaboard of North and South America (Bryan, 1989). An enormous number of flower bulb gene ra and species are found in nature, and they provide material for a wide range of potential ornamental use. The diversity of flower color, form, size, habitat and desirabl e growing conditions of bulbous plants rivals most other forms of vegetation. They can be used in the landscape, as borders and in flower beds, in containers outdoor and indoor and as cut flowers in indoor arrangements. Historically bulbs were introduced in Europe almost 400 years ago (Bryan, 1989). Agapanthus africanus Dur et. Schinz (1893) from South Africa was introduced in England in 1629. Amaryllis bealladona was introduced to Europe in the late 17th century (Traub, 1958a) and the genus Hippeastrum in the beginning of the following century (Tjaden, 1979). Although no registries exist, it is likely that Amaryllis belladonna was originally collected in South Africa at th e time of the spice and slave trade – late 15th century and early 16th century, possibly due to its abundanc e, coastal distribution, floral attributes and commercial value. The slave trade and sugar cane production played a major role in

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2 the distribution of A. belladona around Europe explaining why it is still commonly found in the Canary Islands, Madeira, Spain, Italy, and the Azores, which were main sugar cane growing areas. During the 1500’s, European plant explor ers searched the world recording new species of plants using botanical illustra tions. One of the earliest and best known illustrations in Europe was of Tulipa bononiensis a species of bulb that would become extensively used, appreciated and commercialized. The Dutc h botanist Carolus Clusius, head of the University of Leiden’s Hortus Botanicus, the first botanical garden in Western Europe, received several tulip bul bs and seeds from Turkey and started a collection at the end of the 16th century. Clusius bred the tulips and produced new color variations, but he was mostly interested in its scientific importance and possibly medicinal uses for the bulbs. However, peopl e in Holland were already interested in the flowers as money-makers for the developing ornamental floral trade, and some of Clusius’ tulips were stolen from his gard ens. That was the beginning of the famous “Tulipomania” (Cremers, 1973). Throughout the early 1600’s tuli ps were widely traded in the market and their prices were extremely high. In 1624, one tuli p type was sold for 3000 guilders per bulb, the equivalent of US $1,500.00 nowadays. Since th en the Dutch have bu ilt one of the best organized bulb production and export businesse s in the world. In 2001, over nine billion flower bulbs were produced in Holland, and about 80% were exported. According to the Netherlands Flower Bulb Information Center, the United States is the biggest importer of Dutch bulbs.

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3 Tulips and other spring blooming bulbs ar e still closely associated with the Netherlands; however bulb production is not ex clusive to that area. The world flower bulb industry produces a wide variety of hi gh quality bulbs adapted to many climatic zones that are marketed either as cut flowers, flowering potted plants or landscape plants. Today’s bulb industry is firmly established in areas of the world where the growers can duplicate by various cultural practices the environmental factors required for floral initiation and development. The demand for bu lb crops is high, and the potential exists for greater production and sales of these crops (Johnson et al., 1995). Florida is the second largest floriculture producing state in the US with over $650 million in sales according to USDA, making floriculture a vital part of Florida's agricultural economy. Its hot and humid summers and rare frost occurrences during winter provide an ideal scenario for a varied selection of bulbous plants. The tropical bulb market is practically nonexistent in Florida due to production and market problems. Many tropical bulbs gr ow slowly, some have viral problems, and many consumers are not sure how to grow them However, there is a large selection of bulbs suitable for Florida and other regions in the US in USDA Hardiness Zones 9, 10 and 11 that produce great spring and summer colo r. South Florida's climate is favorable for growing most of the tropical and subtropi cal bulbs. Examples of suitable tropical bulbs for most of Florida include: Af rican Lily, or Lily of the Nile ( Agapanthus africanus ), Amaryllis ( Hippeastrum spp.), Spring Calla Lily ( Zantedeschia spp.), Peruvian Daffodil ( Ismene narcissiflora ), and Rain Lilies ( Habranthus and Zephyranthes ) (Black et al., 1990).

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4 It is important to increase our research concerning flower bulbs, particularly subtropical and tropical bulbs, in order to identify and unders tand cultural factors of as many genera as possible. This will increase th e number of bulbous plants that can be economically produced for the ornamental market. However, to be economically competitive, these plants should be profitable to produce and market, multiply under a wide variety of soil and climatic conditions, adapt to mechanical handling, tolerate air pollution, resist periods of drought, and tole rate low to moderate nutritional status. The present study addressed some of thes e issues and was designed to provide growers with improved methodology and guide lines for the commercial production of two genera of tropical bulbs Habranthus robustus (Herb Lodd. 1831) and Zephyranthes spp. (Herb., 1821). The genera Habranthus and Zephyranthes belong to the Amaryllidaceae family and are known as Rain Lilies since they bloom several times during a season, usually following rainfalls. Li mited information is available in the horticultural literature on cu ltural practices and commerci al production of these two genera (especially of Habranthus) and their ability to adapt and develop under Florida’s climatic conditions. Increased knowledge of these factors could result in the plants becoming important commerc ial crops in Florida. This study was designed to determine the e ffect of environmental factors of light levels, drought stress, and the cultural practi ces of irrigation freque ncy and fertilization on flowering response of Habranthus robustus and Zephyranthes spp. Specific objectives of this study were 1) to pe rform morphological evaluations on Habranthus robustus and Zephyranthes spp. bulbs, comparing those with othe r species of tropica l bulbous plants; 2) to evaluate environmental e ffects on flowering responses for Habranthus robustus and

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5 Zephyranthes spp.; and 3) to develop commercial production and cultural information for Habranthus robustus and Zephyranthes spp.

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6 CHAPTER 2 LITERATURE REVIEW Geophyte, from the Greek geo for earth and phyt on for plant; is a term that refers to plant species with specialized underground storage organs that accumulate food reserves, nutrients and moisture for s easonal growth and developmen t. This group of plants includes both monocotyledonous and dicoty ledonous species (Bryan, 1989) and is collectively referred to as “flowe r bulbs” (Halevy, 1990; Rees, 1985). Geophytes can be separated into four gr oups – true bulbs, corms, tubers and rhizomes. Although they are morphologically di fferent, the undergr ound portions of all types of geophytes perform the same basic f unction – storage. A key factor used to classify geophytes is the precise origin and nature of the tissu e that serves as the primary storage tissue (De He rtogh and Le Nard, 1993). Despite morphological differences, all geophytes share a common characteristic: they have a self-contai ned, highly developed food-storage mechanism. True bulbs have a shortened stem, with a ba sal plate, one or more apical meristems, enclosed flower buds, adventiti ous roots initials, several la yers of fleshy scales and a protective tunic that envelops the bulb. The scales are modified leaves (enlarged leaf bases) and function as the primary storage tissu e in true bulbs. The tunic protects the bulb from drying and mechanical injuries (De Hertogh and Le Nard, 1993). Bulbs can be either tunicate or non-tunicat e depending on the origin of their scales. Concentric layers of scales form tunicate bulbs, such as in Tulipa, Hippeastrum and

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7 Narcissus Non-tunicate bulbs do not develop concentr ic or scaly layers of scales, such as Lilium (Black et al., 1990). Small underground bulbs are either called bulblets or offsets, if they occur at the periphery of the mother bulb. Small aerial bulbs th at occur in either the leaf axils or in the floral parts are called bulbils. Most true bulbs are monocotyledonous such the genera Allium, Amaryllis, Haemanthus, Habranthus, Lilium, Nerine, Tulipa and Zephyranthes (Bryan, 1989). Rhizomes are horizontal, thickened, branch ing storage stems which grow below or along the surface of the soil. Typically, shoots (above ground ) and roots (on the lower surface) arise at right angles from th e cross stem. They are monocotyledonous ( Agapanthus, Canna and Clivia ), and dicotyledonous (some Anemone and Achimenes ). Corms are solid masses of stem tissue with an enlarged basal plate, distinct nodes and internodes, and adventitious root initials, enclosed by several dry, papery scale-like leaves. They are protected against injury and water loss by dry leaf bases that are similar to the tunic that encloses true bulbs. Corms are easily distinguished from true bulbs by the absence of visible storage rings when cr ossly sectioned and by having the basal plate as the primary storage organ. Small corms ar e called cormlets or cormels. Most corms reproduce by annual replacement and are monocotyledonous; examples include Crocus Freesia and Gladiolus (Bryan, 1989). Tubers consist basically of enlarged unde rground stem tissue with a root primodia developing basally and one or more apical shoot meristems (buds) on the underground stem. These buds (also called eyes) arise fr om the nodes and they are arranged in the same spiral pattern characteristic as buds on an aerial stem. Tubers are covered with a

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8 tough skin and do not have scales as in true bulbs. Another distincti on from true bulbs is the absence of a basal plate and protective tunic. They o ccur in both monocotyledonous ( Caladium, Gloriosa and some Zantedeschia ) and dicotyledonous ( Anemone and Eranthis ) genera (Bryan, 1989). Aerial Organs and Tissues Geophytes exhibit a wide variety of flower s, stems and leaves (Bryan, 1989). Their flowers can be single, double, semi-double or multiflowered, depending on the number of petals. According to Du Plessis and Duncan (1989) they can be hysteranthous (leaves appear after flowers), proteran thous (leaves appear before fl owers) or synanthous (leaves and flowers appear simultaneou sly). Floral development can be determinate (number of flowers do not increase after first flower is opened) or indeterminat e (number of flowers can increase after first flower is opened). Stem s can be leafed or l eafless. Plants can be multi-stemmed or single-stemmed. Additionall y, geophytes can be classified as either evergreen or deciduous according to thei r leaf persistence throughout the year. Underground Tissues Three morphological characteri stics are commonly used to describe flowering bulb root systems: branching habit, presence or abse nce of root hairs and contractile root habit. Although most higher plants produce branched root systems (Whittington, 1969), several ornamental flowering bulbs do not. No branch ing has been observed for several genera such as Allium, Crocus, Hyacinthus, Muscari, Narcissus and Tulipa (Kawa and De Hertogh, 1992). It is generally assu med that all plant roots have root hairs, but this is not true for some ornamental flower bulbs such as Crocus flavus (De Munk and De Rooy, 1971).

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9 Contractile roots play a major role in pos itioning bulb crowns and storage organs at a proper soil depth for optimal growth and su rvival of the species. Corms and true bulbs commonly reposition themselves in the ground vi a contractile roots. This mobility of the underground plant organs occurs because of the activity of the contractile roots in response to light and temperat ure (De Hertogh and Le Nard, 1993). Of the bulbs used in their study, Theron and De He rtogh (2001) reported that Hippeastrum, Agapanthus and Zephyranthes have contractile roots, while Scadoxus did not have contractile roots. Habranthus was not used in their study. Rhizomes secure an optimal position by the growing activity of their shoots tips, as reported by Ptz (1998) who found that Hemerocallis fulva rhizomes were able to adjust their own soil depth by elongating the root axis and not via root contraction. Taxonomy and Origin of Geophytes The three most important plant families containing geophytes are Amaryllidaceae, Iridaceae and Liliaceae. Most tropical bulbs, and those used in this study, belong to the family Amaryllidaceae. Bryan (1989) summar ized the origin of many geophytes and stated that they mainly occur between the 23 to 45 North and S outh latitudes, this includes the Mediterranean, South Africa, Middl e East and the Pacific seaboard of North and South America. Geophytes Growth and Development Geophytes must reach a certain physiological stage before they are capable of flowering; this can take less th an a year or as long as six years (De Hertogh and Le Nard, 1993). In several species the abili ty to flower is directly related to the size of the geophytic organ, which varies from species to species. Hippeastrum for example, flowers when bulbs reach a circumference of approximately 20 cm (6 cm in diameter), Scadoxus

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10 flowers when the geophytic organ circumference is 15 cm (4.5 cm in diameter), while Eucharis bulbs need to attain a circumference of 3 to 5 cm (1 cm in diameter) before flowering occurs (Theron and De Hertogh, 2001). A number of factors affect growth, deve lopment, and flowering in bulbs. These include their native habitat and related micr oclimate parameters such as temperature range, rainfall, sun irradiation, photoperiod, al titude, moisture, soil type and nutritional status. With the exception of some equatorial areas th at have fairly uniform environments, geophytes are often exposed to a wide range of climatic conditions during their growth cycle (De Hertogh and Le Nard, 1993). Among environmental factors that affect bulb growth and development, the most important ones are temperature, light and mo isture. These three factors are manipulated to force bulbs, since they act directly on rooting, flower development, shoot elongation, and bulbing (bulb elongation). Temp erature is the major external factor that controls growth, development, flowering, dormancy a nd physiological maturity of bulbs. Light intensity and photoperiod also affect se veral physiological processes, such as photosynthesis, flower aborti on and abscission (Bryan, 1989). Dormancy Bulbs have developed mechanisms to survive seasonal changes in climatic conditions such as low or high temperatur es and drought. Under adverse conditions many bulbs enter a dormant period, in which they do not exhibit any visible external growth. Le Nard (1983) stated that the period of dor mancy mainly corresponde d to the period of bulbing, when the bulb enlarged. However, De Hertogh and Le Nard (1993) defined it as a complex and dynamic physiological, morphol ogical and biochemical state during which

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11 there are no apparent external changes or growth. Internally, however, many physiological and/or morphological events may occur. Roughly, species that go dormant are deci duous as they loose their leaves during adverse periods (winter or s easonal dry periods) and thos e that do not go dormant are evergreen (they keep their leaves all year round). Examples of evergreen bulb genera include Agapanthus, Amaryllis and Clivia Different genera res pond differently to this period of dormancy and have also distinct requirements to break dormancy. Yet, temperature and moisture are the two major factors used to affect bulb dormancy (Le Nard, 1983). Some temperate zone bulb species, such as Tulipa, Daffodil and Hyacinth have cold dormancy requirements; which means they need a critical number of chilling hours in order to bloom the following season (Kamenetsky, 2004). That does not occur in Florida even in the northern areas of the st ate as most winters ar e not sufficiently cold. Consequently, temperate zone bul bs are not used in the lands capes, except as annuals, in these areas. Other species, such as Caladium and Zantedeschia have warm dormancy requirements, which means they need a period of dry and warm temperature in order to bloom next season (De Hertogh and Le Nard, 1993). North and Central Florida’s winters are usually not sufficiently dry or warm fo r some of these species; however, South Florida’s winters are suitable for some of the species. The existence of a rest or dormant peri od is very convenient for horticultural practices because it facilitate s the handling, storage and trans portation of these bulbs. The species used in this study, Habranthus spp. and Zephyranthes spp, go dormant during the cold season and require very little water du ring this period. Howeve r, not all bulb crops

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12 go through a dormant period. Hippeastrum bulbs, for example, do not require a rest period in order to flower, since bulbs can produce infl orescences annually in a greenhouse under conditions of continue d irrigation (Hayne s et. al, 2001). Flowering Process Plants continue to generate new organs even after their embryonic phase, unlike animals. Undifferentiated cells called meristematic cells are responsible for the formation of new organs. The organ produced by the shoot meristem during its post-embryonic phase will depend on the phase of the plant’s life cycle, which can be characterized as: juvenile vegetative phase, adult vegetative phase and reproductive phase. During the juvenile phase, the shoot meri stem initiates stems, true leaves and axilary buds; and during the adult phase, the shoot meristem can form inflorescences which contain sexual organs. In some plants reproduction is the la st of the shoot’s phases; in other plants vegetative growth begins on lateral meristem s when the apex becomes reproductive. In some other plants the apical meristem remains vegetative and lateral meristems generate reproductive structures (Poethig, 1990). The transition from producing one organ to another, which is known as phase change, can be either gradual or abrupt. This process, calle d flower initia tion, marks the end of vegetative growth and is a major determinant of plant reproductive success (Poethig 1990). The flowering process involves five successive stages: induction, initiation, organogenesis, maturation (growth of the flor al parts), and anthesis (De Hertogh and Le Nard, 1993). Important aspects of this proce ss are: 1) the signal s received by the plant that instigate the proces s, 2) their transport to the shoot apex, and 3) the changes in the shoot apex during floral di fferentiation (Evans, 1993).

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13 The successive steps of the flowering process are controlled by several factors and each takes place in a determined period of the growth cycle. Controlling these processes in bulbous plants, as well as other horticultu ral crops, can promote or retard flowering, prevent flowering or induce flower aborti on (De Hertogh and Le Nard, 1993). Promoting or retarding flowering permits out of s eason and/or year-round commercial flower production. Preventing flowering is necessary for the producti on of some bulbs, such as Iris hollandica Inducing flower abortion can prom ote bulb development in some bulb species such as Tulipa and Hippeastrum The flowering process can be controlled by applying specific treatments (temperature, mois ture, light, or plant gr owth regulators) to the bulb. However, precise knowledge of bulb peri odicity is essential to control flowering (Bryan, 1989). Flower initiation takes place at different times of the year and at different stages in the development of bulbs. Seven different type s of flower initiation have been identified in commercially grown bulbous plants (Hartsema, 1961): 1. Flowers are formed during spring or early summer of the year preceding the one in which they flower ( Narcissus, Galanthus, Leucojun and Convallaria ). 2. Flowers are formed after the end of the assimilation period ( Crocus, Hyacinthus, Iris reticulata and Tulipa ). 3. Flowers are formed some time after new grow th matures, in winter or early spring (most Iris spp). 4. Flowers are formed during the storage period and complete development after planting ( Begonia tuberosa, Dahlia and Lilium ). 5. Flowers are formed afte r replanting in spring ( Galdiolus Anemone and Freesia ). 6. Flowers are formed more than a year before flowering ( Amaryllis belladonna and Nerine sarniensis ).

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14 7. Flower formation occurs alternatively with leaf formation during the whole assimilation period ( Zephyranthes and Habranthus ). In this case, both young developing flower buds and one year-old flower buds are present in the bulb. Anatomy and Physiology of Flower Initiation All the aerial parts of a plant (excludi ng the cotyledons) are produced by the shoot apical meristem. This meristem is form ed during embryogenesis and comprises a group of undifferentiated cells that generate differe nt organs and structur es such as stems, leaves and flower. The root system of a plan t is produced by the root meristem (Poethig 1990). The shoot apex meristem at a certain poi nt in the growth cy cle undergoes a phase change, caused by floral induc tion in response to exogenous (such as daylength, nutrient status, or temperature) and endogenous (related to internal factors su ch as plant age and metabolic status) factors, and becomes re productive producing flow er buds instead of leaves. This transition is not irreversible in plants and in some species under certain environmental conditions, leafy shoot forma tion occurs after flower formation in a phenomenon known as inflorescence reversion (Pouteau et al. 1997). Three different models have attempted to describe the control of flowering in plants. The first model is known as the “flo rigen concept” (Evans, 1971) which suggested that substances or signals are transferre d across grafts between reproductive “donor” shoots and vegetative “recipients”, and that a flower-promoting hormone called florigen transports them to the shoot apex via the phloem. Chailahjan (1937) suggested that the florigen hormone was produced in leaves under favorable photoperi odic conditions. This model could not be proved, which lead to a second model called nutrient diversion hypothesis. This suggested that external condi tions could raise the amount of assimilates moved to the apical meristem, which were responsible for flower induction (Bernier,

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15 1988). A third model, called “multifactorial control model”, proposed that several promoters and inhibitors (including phytohor mones and assimilates) were involved in flowering induction; and that flowering woul d only occur if these were present in the apical meristem in proper concentrations and specific time in tervals (Evans, 1995). Genetic investigations of flowering in se veral species support the assumption that several factors act toge ther to control flowering. Most of these studies have been done in Arabidopsis and have provided recent advances in better understanding of this process (Guo et al, 1998; Koornneef et al., 1998; Simpson, 2002). A number of genes are involved in flower initiation, such as the flowering-time, which integrates the signals and act as pr omoters or repressors of flowering, the meristem-identity, which determines the fate of newly formed primor dia (shoot/leaf-, or a flower-primordium), and the organ-identit y, which directs the formation of various flower parts (reviewed in Ya nofsky, 1995). Peeters and Koornn eef (1996) have identified several genes that control th e timing of the transition from leaf production to flower production by mutant analysis. Th ere is still much to be learned about the functions of these different genes, however, it has been s uggested that some of these flowering-time genes are involved in the part itioning or metabolism/sensi ng of compounds that might play a part in the endogenous plant signal(s), such as the plant hormone gibberellin, and sucrose concentrations (Nilsson et al. 1998). There is a small group of plants that can be induced to flower artificially by the application of chemicals. Manochai (2005) test ed the application of potassium chlorate (KClO3) to either the root or the foliage of Dimocarpus longan and obtained successful results at all times of the year.

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16 During flower induction and early flower development the apical meristem undergoes several morphological alterations (T ooke and Battey, 2003). Changes in local forces at the surface of the shoot apex have been proposed to play an important role in organogenesis (Fleming et al., 1997), which is the step that follow s initiation in the flowering process. Albrechtov et al. (2004) examined the role of these local forces during photoperiodic flower induction in Chenopodium rubrum by measuring the changes in shape of the apical dome, and changes in cell wall properties. The results obtained confirmed that early changes at the surface of the apical meristem affect the process of floral transition. The relative size of the apical meristem has also been proposed as an important factor in the de velopmental (vegetativ e/reproductive) switch. After a signal is received by the plant and transferred to the shoot apex, the meristematic cells begin the process of orga nogenesis, the formation of tissues by cell differentiation. These tissues mature forming the floral organs. Different species have different mechanisms with which they contro l the development of their flower organs (stamens, carpels, etc). Most species deve lop their carpels and stamens simultaneously, but in some species the male flower orga ns grow first (these are referred to as protandrous flowers) and in ot hers the female organs grow first (and these are referred to as protogynous flowers). A few species like pumpkin ( Cucurbita pepo ) develop male flowers first, next hermaphroditic ones, fi nally female and at last parthenocarpous flowers (lacking an embryo) that produce s eeds agamically, not involving the fusion of male and female gametes, as occurs in sexual reproduction (Evans, 1993). A flower is considered a modified stem with shortened in ternodes and bearing modified leaves at the nodes. In essence, a flow er is structured as a modified shoot or axis

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17 with an apical meristem that does not continue to grow. A typical flower contains a pedicel (stem) with a torus or receptacle at the end. Th e four main parts (or whorls) of a flower sit on the receptacle, and they are: the calyx (group of sepals); co rolla (group of petals); the androecium (male organs – filaments and anthers); and the gynoecium (female organs – stigma, style, ovary and ovul e). Plant species show a wide variety of modifications from this plan (Eames, 1961). The basic function of flowers is to mediat e the union of male and female gametes, in the process called pollination. The period of time during which the flower is fully expanded, functional, and thus allow the pol lination process can take place is called anthesis, and it can vary from few hours to weeks. During the maturation phase, most assimilates obtained by the plant are used to develop the floral parts. As soon as the flower is fully expanded and opened, assimilate s start to be mobilized and used for seed development, and that is when the process of flower senescence be gins (Poethig, 1990). Factors Affecting Flower Initiation In some species, the timing of flowering is primarily influenced by environmental factors, which determine the time of year an d/or growth conditions that are favorable for sexual reproduction and seed maturation. Thes e factors include photoperiod (day length), light quality (wave length), light quan tity (photon flux density), temperature, vernalization (exposure to a de fined number of hours below a critical temperature), and nutrient and water ava ilability (Halevy, 1990). Some species are not very sensitive to e nvironmental variables and these appear to flower in response to endogenous factors such as plant size (bulbs ar e an example of that) or number of vegetative nodes. Flowering can al so be induced by stresses such as nutrient deficiency, drought, and overcrowding. This re sponse enables the plant to produce seeds,

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18 which lay dormant until the return of favor able environmental f actors (Levy and Dean, 1990). Photoperiod Photoperiod is defined as the length of time a plant is exposed to light and/or darkness within a 24-hour period. Flower induc tion, initiation and/or development of many plant species are greatly affected by photoperiod and can be synchronized during the year by manipulating night length (Garne r and Allard, 1920). According to Thomas and Vince-Prue (1997), plants with photoperiodic flowering responses are divided into five groups according to the amount of light/dar kness they require: s hort-day (SD) plants, long-day (LD) plants, day-neutral (DN) plants, intermediate-day plants, and ambiphotoperiodic-day plants. Shortand longday plants can also be subdivided into: obligate or qualitative, and facultative or quantitative. Photoperiodi c responses are often interspecific and can vary within cultivar s of a species (Martson and Erwin, 2005). In most plants the amount of darkness is what determines initiation; a night break (or addition of extra light during the dark period) can inhibit or promote flowering (Evans 1993). Rees (1985) reported that Hippeastrum bulbs have no response to photoperiod regarding flowering. However, De Hert ogh and Gallitano (2000) determined that photoperiod, as well as temperatur e, affect leaf size in the ‘Apple Blossom’ cultivar of Hippeastrum Light quality and quantity There are two different plant behaviors rela ted to light quality: light dominance and dark dominance. Light dominants are plants that are susceptible to changes in the spectral

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19 distribution during the light pe riod. Dark dominants are plants that need uninterrupted dark periods in order to flow er (Fankhauser and Chory, 1997). Flower initiation and time to flower in LD plants are related to light quality and to the timing of light treatments in the photope riod (Thomas and Vince-Prue, 1997). Pringer and Cathey (1960) examined the effects of different types of light on Petunia flowering and concluded that they flower 2 to 3 week s earlier if exposed to incandescent light (containing red and far-red light) than when exposed to fluorescent light (which has no far-red light). Red light appears to be effective earlier in the photoperiod for LD plants, while farred light seems to be important in the flow er inductive response later in the photoperiod for LD plants, which according to Thomas a nd Vince-Prue (1997) is different from SD plants. Studies made by Vince (1965) in Lolium temulentum showed that red light used for 8 hr at the beginning or at the end of the photoperiod induc ed flowering, but when it was used at the beginning it promotes stronger flowering. Temperature In bulbous plants, temperature is the majo r external factor controlling growth, development and flowering (promoting or de laying it). Temperatur e also affects bulb dormancy in some genera and the physiol ogical maturity of the bulbs (Bryan, 1989). Doorduin and Verkerke (2002) investigat ed bud development and flowering of Hippeastrum under 15 to 25oC and observed that at higher te mperatures larger bulbs with more leaves developed, but the percen tage of bulb dry matter decreased. In Scadoxus temperature affects not only flowering but also bulb development. The optimal temperature to overcome dormancy of Scadoxus bulbs is 10 to 15oC, which is low enough to injure several ot her species (Bryan, 1989).

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20 Flower formation does not occur in Amaryllis belladonna at 9o, 13o or 31oC, and the optimal temperature for flower initiation on this species appears to be 17oC. However, for non-planted Amaryllis belladonna bulbs, the optimal temperature for flower formation is 23oC (reviewed in Theron and De Hortogh, 2001). Mori and Sakanishi (1989) demonstrated that for Agapanthus flowering occurred when plants were continuously grown at 10 o or 15oC; in contrast, flowering was inhibited when plants were grown at 20oC or higher from September onward. They also found that flowering could be accelerated by gr owing plants at a minimum of 20oC after the end of November in combination with a 16-hour photoperiod. Hartsema (1961) reported that flower formation of Zephyranthes rosea occurred at 13o to 18oC and flowering occured at 22oC, with soil moisture being important. No flowering occurred at 30oC. In Zephyranthes candida inflorescences were formed in spring and when plants with at least two previously formed inflorescences were subjected to 10o, 15o, 23o or 30oC in October, earliest flowering occurred on plants exposed to 23oC (Hartsema, 1961). Vernalization Vernalization is a period of cold temperat ure treatments that accelerates flowering in some plant species. Many biennial (two -year) plants require a temperature below a critical level for a definite time period before flowering can occur. Plants can either have an obligate or quantitative ve rnalization period. Obligate vern alization refers to plants that require cold temperatures for a period of time in order to flower. Quantitative vernalization refers to plants that do not require cold temperat ures to flower but start their flowering period earlier under co ld temperatures. Additionally, there are plants that do not respond to cold temperatures (reviewed in Yan and Wallace, 1995).

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21 Flower Initiation Process in Bulbous Plants Most true bulbs have a sympodial branch ing system (superposed branches) and at flower initiation a growing point is formed laterally in the apex. This growing point develops a certain number of leaves before an inflorescence develops, alternating flower and leaf formation throughout the entir e growth period (Hartsema, 1961). The flower/inflorescence emergence is delayed comp ared to leaf emergence thus leaves and flower/inflorescence above ground have a diffe rence of one generation between them, and the inflorescence appears to be latera l to leaves (De Hertogh and Le Nard, 1993). Blaaum (1931) described the general flower ing process in true bulbs and according to his studies bulbs undergo the following st ages: I) the meristem is vegetative and produces leaves; II) the last leaf and new growing point are formed; III) a certain number of leaves are formed; IV) then the flower /inflorescence meristem is formed; followed by flower and floral parts formation. More re cently, De Munk and Van der Hulst (cited in Theron and De Hertogh, 2001) ha ve described the flower ing process as: Stage I) vegetative; Stage II) formation of spathe; Stag e III) beginning of flower initiation; Stage IV) flower development and anthesis; Stage V) flower senescence and vegetative growth. Theron and De Hertogh (2001) showed that Hippeastrum bulbs initiate the flowering process in the spring, the differen tiation period lasts from 18 to 24 months and anthesis occurs in the spring. Flower Bulb Cultivation Field production of bulbous crops occurs where soil and climate have advantageous characteristics. A large bulb industry deve loped in the Northwest region of the United States because soils usually do not freeze enough to damage the bulbs and abundant rainfalls create favorable growing conditions for them. Mild temperatures and abundant

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22 moisture favor the production of bulbs havi ng a multi-year producti on cycle. In more severe climates, such as the Midwest and Nort heast, tender bulbs must be removed from the soil in the fall and stored dur ing the winter (Johson et al., 1995). Some bulbs grow well in light sandy or gr avelly-type soils. However, most bulbs grow best in loams with high organic matte r content. Generally, bulbs do not grow well in water-logged or heavy cl ay soils (Johson et al., 1995). The majority of bulbous crops are planted in the early fall; howe ver, some varieties are planted during the spring, such as Gladiolus, Gloxinia and Begonia The bulblets remain in the ground for 1 to 3 years until they reach harvestable size. Some tender species require “lifting” or removal from th e soil in the fall to avoid freeze damage (Johson et al., 1995). The harvesting process is usually done mechanically, using methods similar to those used in onion production. Bulbs are lift ed from the soil and deposited onto a beltconveyor that moves them into the harveste r, which shakes the bulbs to loosen and separate the soil. After harvesting, bulbs are sorted, graded, and damaged bulbs are discarded (Johson et al., 1995). Flower bulbs present several different gr owth habits and when cultivated under growing conditions much different than their na tive habitat, they may drastically change their growth and floweri ng habit. For example, Ornithogalum which is a deciduous perennial, becomes an evergreen when grown in the tropics, does not go dormant, blooms constantly, and does not produce bulbs (Halevy, 1990). Flower Bulb Forcing Forcing is defined as the regulation of bulb growth and development under greenhouse controlled environmental conditions (De Hertogh, 1977). Tr ue bulbs, as well

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23 as corms, rhizomes, and tubers can be for ced. According to De Hertogh (1977), in order to succeed using this process a grower must fully understand: a) the origin and morphology of the species, b) the production and growth cycle of the species, and c) the influence of various environmental fact ors on the development of the species. The complete forcing system has been di vided into four distinct phases: I) production, II) programming, III) greenhouse and IV) marketing. The exact programming of temperatures and times varies from species to species and might not be applicable to all cultivars of all sp ecies. The complete process shoul d be based on marketing time and bulbs should finish the process at proper stage of development (De Hertogh, 1974). The basic bulb production cy cle initiates when bulbs are harvested, sorted and graded (according to their circumference meas urements). The process continues with the storage of bulbs in warm temperatures to full y develop the floral or gans. In the fall, the bulbs are planted, kept moist and under temp eratures low enough to promote flowering and bulbing. In spring, the flor al stalk elongates and the pl ant flowers. Some species require the removal of flowers, to increase bulb size (De Hertogh, 1977). Successful forcing of flowering bulbs in a greenhouse is based on seven factors: temperature (the most important), watering, li ght, fertilization, vent ilation, sanitation, and pest control. Forcing can be either acceler ated or delayed by mani pulating these factors (De Hertogh, 1996). Hippeastrum bulbs can be forced for either fr esh cut flowers or flowering potted plants. The key factors for forcing Hippeastrum bulbs are: 1) use bul bs larger than 20 cm in circumference (6 cm in diameter), 2) rem ove the bottom half of the roots, 3) cure the bulbs for 10 days at 17o to 23oC, 4) store the bulbs at 9o to 13oC for at least 8 weeks

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24 before planting, 5) package bulbs to avoid r oots drying out if transported, 6) plant bulbs in well-drained planting medi um, 7) use bottom heat of 22oC in the greenhouse (reviewed in Theron and De Hertogh, 2001). Tropical Bulbs and Amaryllidaceae Most tropical bulbs, and in pa rticular those used in this study, belong to the family Amaryllidaceae, which is a very diverse family with species on almost all continents and under various climatic conditions (Meerow and Snijman, 1998). Amaryllidaceae classification includes the following: Kingdom: Plantae Subkingdom: Tracheobion ta (Vascular plants) Superdivision: Spermatophyta (Seed plants) Division: Magnoliophypa (Flowering plants) Class: Liliopsida (Monocodiledoneous) Subclass: Liliidae Order: Liliales / Amaryllidales / Asparagales Family: Amaryllidaceae According to Watson and Dallwitz (1992) the family Amaryllidaceae comprises approximately 60 genera and 800 species. Some of the important horticultural crops are Amaryllis, Clivia, Crinum, Eucharis, Haemanthus, Hippeastrum, Hymenocallis, Lycoris, Narcissus, Nerine, Scadoxus and Zephyranthes The two most recent formal classifications of the Amaryllidaceae family are those of Mller-Doblies and Mlle r-Doblies (1996) and Meerow and Snijiman (1998). Hyacinthaceae, is also considered a rela ted family, however Agavaceae, Hypoxidaceae,

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25 Haemodoraceae or Alstroemeriaceae with which Amaryllidaceae is sometimes associated, are not considered related families. Plants in the Amaryllidaceae family have distinct habits, however most of them are perennial, bulbaceous and have contractile roots. Leaves are mostly deciduous and simple, with entire lamina, entire margins and parallel venation. Flowers may be solitary or produced on different types of inflorescences. Bulbous plan ts in this family have thickened underground storage organs whic h enable them to survive unfavorable environmental conditions and may also f unction as propagative units (reviewed by Theron and De Hertogh, 2001). Halevy (1990) demonstrated that when flower bulbs are produced under growing conditions that are dissimilar to their indige nous environments, their growth habit can be altered. Therefore, it is importa nt to understand the effects of temperature, light, nutrition, growth regulators, and other environmental factors on bulb growth, development and the flowering process, to significantly expand the horticultural usage of the Amaryllidaceae family (Theron and De Hertogh, 2001). Hippeastrum spp. The name Hippeastrum comes from the Greek hippe us, meaning knight and astron meaning star. Hippeastrum is an important genus of the Amaryllidaceae family which comprises about 70 species, such as H. argentinum, H. aulicum, H. barbatum, H. correiense, H. elegans, H. evansiae, H. leopoldii, H. miniatum, H. morelianum, H. pardinum, H. psittacinum, H. puniceum, H. reginae, H. reticulatum, H. rutilum, H. stylosum, H. vittatum and H. reticulatum (Rees, 1985) There are more than 300 cultivars and most of the horticulturally important ones were bred by Ludwig, Warmenhove and

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26 van Meeuven in the Netherlands and by HADE CO (Barnhoorn) in S outh Africa (Read, 2004). Hippeastrum can be considered a tropical plant by origin as it is indigenous to Central and South America, being centered in Brazil and Peru, and distributed from Mexico to Argentina (De Hertogh and Le Nard, 1993). Hippeastrum is commercially known as Amaryllis, however, the true Amaryllis ( A. belladonna ) originated in South Africa and it is not widely used. There are se veral differences between them such as the presence of a solid scape and the absen ce of scales between the filaments in A. belladonna (Goldblatt, 1984). Hippeastrum spp has large and showy flowers with many bright colors (red, pink, orange, white or bi-colored) consisting of several flower type s: trumpet-flowered, belladonna types, reginae type s, leopoldii types, mini atures, doubles, and orchidflowered. They are grown mainly as potted plants or as cut flowers, but they can also be grown in the landscape in subtropica l and tropical areas (Schulz, 1954). The bulb has a sympodial branching system, which means that the terminal bud dies or ends in an inflorescence, and growth of sympodial shoots continues from lateral buds. At flowering initiation, a la teral growing point is formed on the side of the apex. It develops in a sequence of four leaves and an inflorescence. Bulblets are initiated in the axils of senescing bulb scales in the outer parts of the bulb and they produce nine leaves before initiating the first inflorescen ce (reviewed in Theron and De Hertogh, 2001). Scadoxus multiflorus – Blood Lily The genus Scadoxus contains 9 species: S. cinnabarinus, S. cyrtanthiflorus, S. longifolius, S. membranaceus, S. multiflorus, S. nutans, S. pole-evansii, S. pseudocalus and S. puniceus (Friis and Nordal, 1976). Scadoxus is closely related to Haemanthus ;

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27 they were treated as a single genus at one time but were divided by Friis and Nordal (1976). The major differences between Scadoxus and Haemanthus are related to their geophytic organs, growth habits, and number and shape of their leaves (Snijman, 1984). Scadoxus multiflorus is one of the most horticultur ally relevant species in the genus, its name derived from doxus meaning glory or splendor, and multiflorus referring to many flowers (Jackson, 1990). Scadoxus multiflorus is endemic to s outhern and central Africa and was established by Rafinesque (183 6). This species incl udes three subspecies: katherinae longitubus and multiflorus and the major differences between the subspecies are height of the plants, lengt h of the perianth tubes and the extent of the perianth segments (Du Plessis and Duncan, 1989). Agapanthus africanus – African Lily The genus Agapanthus was established by L’He ritier (1788). The name Agapanthus is derived from the Greek agap that means love and anthos that means flower (Snoeijer, 2004). It was placed in the Liliaceae family, later moved to the Amaryllidaceae family. Agapanthus is endemic to southern Africa and the first species collected Agapanthus africanus was described in 1679 by the name Hyacinthus africanus tuberosus which is sometimes still referred to as Agapanthus umbellatus (Leighton, 1965). Agapanthus is a variable genus, all species are rhizomatous with similar appearances. Botanists consider it difficult to cl assify them into distinct species. Plessis and Duncan (1989) identified about 10 species indigenous to Southern Africa. Agapanthus africanus is a summer-flowering evergreen species, with its perennial geophytic organ a rhizomatous rootstock with c ontractile roots. Flowers are formed in an umbel inflorescence on a leafless scape. Mo ri and Sakanishi (1989) observed that A.

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28 africanus meristem was vegetative in October, fl ower initiation began in November and flower differentiation occurred in December. Habranthus robustus and Zephyranthes spp. – Rain Lilies Habranthus robustus was established by Herber t and Loddiges (1831), and Zephyranthes spp. was established by Herbert (1821). The name Habranthus comes from the Greek habros, meaning graceful and anthos meaning flower, and Zephyranthes comes from the Greek, zephyros meaning "the west wind" and anthos, meaning flower. These two groups of summer-flowering small bulbs of the Amaryllidaceae family are native to the southeastern United States, Central and South America (Hume, 1935; Traub, 1958 b). The genus Habranthus contains about 40 species, and among the horticulturally important are H. tubispathus and H. robustus which is native to Brazil (Read, 2004). The Zephyranthes genus contains about 60 species and the most horticulturally important are Z. candida Herb and Z. grandiflora Lindl, but other commercial ly grown species include: Z. andersonii Baker, Z. atamasco (L.) Herb, Z. drummondii Z. primulina T.M. Howard & S. Ogden, Z. rosea Lindl, and Z. citrina Baker. There are several named cultivars derived from interspecific crosses (Van Scheepen, 1991). Habranthus and Zephyranthes geophytic organs are pere nnial bulbs covered with a dark tunica, contractile roots, deciduous leaves with sheath ing basis and linear blades, and isolated flowers (The ron and De Hertogh, 2001). Habranthus robustus flowers are very similar to those of Zephyranthes and Habranthus have at times been included in the genus Zephyranthes because of that. Both are commonly called “rain lilies” because of their tendency to bloom after rain periods (Fellers, 1996).

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29 Habranthus robustus is often called Zephyranthes robusta incorrectly, which adds to the confusion between it and Zephyranthes grandiflora The very subtle differences between genera are based on spathe characters position of flowers, symmetry of corolla, insertion of anther filaments, number and le ngth of filaments, and number of seeds per locule in capsule (Fernnd ez-Alonso & Groenendijk, 2004). Zephyranthes flowers point straight up and ha ve equal lengths stamens, while Habranthus flowers point upward but at an angle and have stamens of different lengths; these characteristics are commonly used to separate the two genera. Additionally, Habranthus flowers tend to have zygomorphic fl owers (bilaterally symmetrical), while Zephyranthes are actinomorphic flowers (radially symmetrical). The flowers of Habranthus are clearly distinguished by filament s of 4 different lengths and always longer than the perianth tube. Zephyranthes flowers are declinate and distinguished by filaments of two very similar lengths (Fernndez-Alonso & Groenendijk, 2004). Zephyranthes and Habranthus can also be distinguished by the shape of their seed. Zephyranthes seeds tend to be more D-shaped or wedge shaped while those of Habranthus are more openly obliquely winged. These two genera are al so distinguished phylogentically as, Zephyranthes apparently have 2-3 diffe rent origins according to nrDNA spacer sequences (Meerow et al, 2000). The differences between Habranthus and Zephyranthes are not readily apparent to most growers and consumers but the two gr oups of bulbs seem to have different flowering periods and re-blooming characte ristics. The results of this study will contribute to a better understan ding of these species and their cultural differences. This

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30 study will also help growers determine which sp ecies grow best at different times of the year and maximize seasonal sales and facilitate their commercialization.

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31 CHAPTER 3 BULB MORPHOLOGY Geophytes were defined by Ra unkier (1934) as plant sp ecies with specialized underground storage organs that accumulate food reserves, nutrients and moisture for seasonal growth and development. They ar e usually collectively called “flower bulbs” and can be separated into four groups: true bulbs (tunicate and nontunicate), corms, tubers and rhizomes. Although morphologically different, all types of geophytes perform the same basic function, st orage of photosynthates. Most geophytes have a shortened stem, a basa l plate, one or more apical meristems, enclosed flower buds, adventiti ous root initials, se veral layers of fleshy scales (modified leaves), and a protective t unic. The size of storage organs vary tremendously among species (Proches et al., 2006), as well as their leaf arra ngements and flower formation. Some genera, including Tulipa and Narcissus (Caldwell and Wallace, 1955) have been extensively examined. However, only limited in formation is available in the literature concerning morphology of ot her tropical bulbs. This portion of the present study was designed to compare and contrast the morphology and flower formation of Hippeastrum hybridum, Scadoxus multiflorus and Agapanthus africanus and compare them to Habranthus robustus and Zephyranthes spp, which were species used in the subsequent phases of this study. Comparative Study Materials and Methods : The five species of tropical bulbous plants listed on table 3-1 were grown under same conditions, dissected by freehand sections, stained with

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32 Safranin, and examined during mature and im mature stages of development. Digital images and drawings were produced to de termine similarities and differences in morphology and floral development of the five species examined. Table 3-1 – Tropical geophytes used in the morphological studies: Species Common Name Family Source Hippeastrum hybridum Amaryllis Amaryllidaceae Agristarts, Inc. Scadoxus multiflorus Blood Lily Amaryllidaceae Agristarts, Inc. Agapanthus africanus Agapanthus Alliaceae Agristarts, Inc. Habranthus robustus Rain Lily Amaryllidaceae UF stock Zephyranthes spp. Rain Lily Amaryllidaceae UF stock Seventy two plugs of Hippeastrum hybridum, Scadoxus multiflorus and Agapanthus africanus were obtained from Agristarts Inc. in Apopka, Florida during the first week of March 2004. Habranthus robustus and Zephyranthes hybrids bulbs were obtained from University of Florida stock where they had been grown in ground beds for a year prior to the beginning of this study. The Zephyranthes hybrids used in this study were: Z ‘Paul Niemi’, Z ‘Jo Ann’s Trial’ and Z ‘Fadjar’s Pink’, which were hybridized by Fadjar Marta in Jakarta, Indonesia. All geophyte plugs and Habranthus and Zephyranthes bulbs were transplanted into 15 cm plastic pots using sphagnum peat ba sed Fafard No. 2 soiless growing medium (Agawam, MA) consisting of 70% Canadi an sphagnum peat, 10% perlite and 20% vermiculite. Plants were placed in a gree nhouse (which provided 11% shade), natural photoperiod and a temperature range of 31/24oC (day/night). From planting through establishment all pots were irrigated ever y other day with 250ml of water, except Hippeastrum which was watered daily with same amount of water. After establishment (first week of April), plants were fertigated twice a week with 250ml of water containing

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33 Peters Professional ‘Florida Special’ water soluble fertilizer 20N-4.7P-16.6K (Scotts Co., Marysville, OH) with N at 150 mg L-1. Plan ts were grown in the greenhouse for ten months. From June to November 2005, a series of fr eehand sections of the four species of bulbs and Agapanthus rhizomes were made. The sections were mounted in glycerin and stained with Safranin for 24 hours to faci litate the observation of specific tissue components. Sections were examined with a Wild M5 dissecting scope and light microscopy images were obtained with a Ze iss Tessovar with an Rts Contax camera attachment. Freehand sections of the meristematic re gions of the five species studied were mounted in glycerin (but not stained) and light microscopy images were obtained with a Nikon Labophot-2 microscope and a Nikon E 4500 digital camera attached to it. Drawings, based on observations of numer ous freehand sections, were made to demonstrate the general anatomy and morphol ogy of each of the five species examined. Results and Discussion Bulb type and size : Observations and measurements made in this study revealed that Zephyranthes and Habranthus bulbs were similar in shape but that Zephyranthes bulbs were smaller than Habranthus bulbs. Mature Zephyranthes bulbs were approximately 6 cm tall with a diam eter of about 6 cm, while mature Habranthus bulbs were approximately 8 cm tall with a diamet er of about 8 cm. Both bulb genera were covered with a fine, papery dark brown tunica. Hippeastrum bulbs were covered with a fine, papery dark brown tunica and were similar to Zephyranthes and Habranthus in shape but were larger, as mature bulbs were approximately 12 cm tall with a diameter of about 12 cm (Table 3-2 and Figure 3-1). Zephyranthes, Habranthus and Hippeastrum

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34 form true tunicate bulbs, with concentric layers of modified leaves called scales (Figures 3-2, 3-3 and 3-4). Scadoxus geophytic organs were true bulbs form ed by concentric layers of scales (approximately 6 cm tall when mature) with a thick (4 cm tall) rhizomatous structure at their base and covered by a fine, papery dark brown tunica (Figure 3-5). Scales of Scadoxus were thicker (0.5 cm) than scales of Zephyranthes, Habranthus and Hippeastrum which averaged about 0.2 cm in thickness. Agapanthus geophytic organs were rhizomes and did not have layers, scales or a tunica as did the other four previous studied species (Figure 3-6). A mature Agapanthus rhizome was approximately 20 cm long with a diameter of approximately 10 cm (Table 3-2). Table 3-2 – Geophyte size and charac teristics of plants examined Genera Geophytic organ Height (cm) Diameter (cm) Scales present Tunica present Hippeastrum True bulb 12 12 Yes Yes Habranthus True bulb 8 8 Yes Yes Zephyranthes True bulb 6 6 Yes Yes Scadoxus Bulb/rhizome 10 12 Yes Yes Agapanthus Rhizome 10 No No

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35 Figure 3-1: From left to right Zephyranthes, Habranthus, Hippeastrum Scadoxus bulbs, and immature Agapanthus rhizome. Bar = 10 cm. (A) (B) Figure 3-2: Sections of Zephyranthes bulbs. Longitudinal section (A) and photomicrograph of a cross section (B). Bar =1 cm.

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36 (A) (B) Figure 3-3: Sections of Habranthus bulbs. Longitudinal section (A) and photomicrograph of a cross section (B). Bar = 1 cm. (A) (B) Figure 3-4: Sections of Hippeastrum bulbs. Longitudinal section (A) and photomicrograph of a cross section (B). Bar = 1 cm. (A) (B) Figure 3-5: Sections of Scadoxus bulbs Longitudinal section (A) and photomicrograph of a cross section (B). Bar = 1 cm.

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37 (A) (B) Figure 3-6: Longitudina l sections of an Agapanthus rhizome. Bar = 1 cm. Leaf arrangement and morphology Hippeastrum, Habranthus and Zephyranthes bulbs had similar leaf arrangements and leaf morphologies. Leaves were perennial, basal, simple, linear, glabrous, with entire margins and parallel venation. They emerged from the center of the bulb, with leaves two-ranked as the blade of each new leaf emerged 180o from the previous leaf, thus older leaves were always on the outside and younger leaves on the inside of the leaf cluster (Figure 3-7, 3-8 and 3-9). Each leaf was composed of a photosynthetic leaf blade and a non-photosynthetic storage leaf base (scale) which thickened during the growth cycle forming true bulbs. Jones and Emsweller (1936) stated that each new leaf of Allium cepa developed on the side of the apical meristem opposite to the preceding blade by an upward growth of tissue surrounding the apical meristem. We observed that Hippeastrum Habranthus and Zephyranthes leaf production occurred in a simila r process. According to Mann (1960) A. cepa bulbs contain leaves that are morphol ogically distinct from each other, and

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38 Kamenetsky (1994) classified them according to their function: protective, storage and assimilation. Black et al. (1990) described Hippeastrum leaf blades as being up to 60 cm long and up to 5 cm wide. We observed that Habranthus leaf blades were 10 to 20 cm long and 1.5 to 2 cm wide; while Zephyranthes blades were 10 to 20 cm long and 1 cm wide. As part of this study, a determination of nu mber of leaves produced weekly for a period of one year of Habranthus and Zephyranthes bulbs was made (Figures 3-41 and 3-42). Agapanthus rhizomes have simple, linear, pere nnial glabrous leaves with entire margins and parallel venation. Leaf blades were up to 50 cm long and approximately 4 cm wide. Leaf arrangement was alternate but we observed that the l eaves did not emerge from the center as they did in Hippeastrum, Habranthus and Zephyranthes, but from the lateral sides of the rhizome (one at a time). T hus, the older leaves were external to inner new leaves. Another distinction between Agapanthus and Hippeastrum, Habranthus and Zephyranthes bulbs was that leaf base s did not thicken and form scales, but remained thin and papery (Figure 3-10 C). Scadoxus leaves were shorter, wider and thinner than leaves of Hippeastrum, Habranthus, Zephyranthes and Agapanthus and they had a distinct oblong shape, prominent mid-ribs, undulating leaf margins, an d leaf blades were 30 to 40 cm long and 9 to 13 cm wide. Scadoxus leaf arrangement was distinct fr om the previous four species. The first two leaves emerged singularly and we re distinctly differe nt from subsequent leaves. After the first two leaves senesced, a cl uster of leaves arose from the center of the bulb all at once. They were held together by a structure called a pseudo-stem (a false stem formed by the sheathing and overlapp ing of the leaf bases – Figure3-11 C).

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39 Table 3-3 – Leaf size of plants examined Species Leaf blade length (cm) Leaf blade width (cm) Hippeastrum Up to 60 5 Habranthus 10 to 20 1.5 to 2 Zephyranthes 10 to 20 1 Scadoxus 30 to 40 9 to 13 Agapanthus Up to 50 4 (A) (B) (C) Figure 3-7: Hippeastrum leaf arrangement. From top (A ), side (B) and individual leaf with continuous leaf base / scale (C). Bar = 1 cm. (A) (B) Figure 3-8: Habranthus leaf arrangement. From top (A) and side (B). Bar = 1 cm.

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40 (A) (B) Figure 3-9: Zephyranthes leaf arrangement. From top (A) and side (B). Bar = 1 cm. (A) (B) (C) Figure 3-10: Agapanthus leaf arrangement. Overlapping of leaf bases (A), new leaves emerging from lateral side of the rhiz ome (B), and individual leaf with continuous papery leaf base (C). Bar = 1 cm. (A) (B) (C) Figure 3-11: Scadoxus leaf arrangement. First emerged leaf (A), photomicrograph of a longitudinal section showing leaf sheat hing in a pseudo-stem (B), and a group of mature leaves in a pseudo-stem (C). Bar = 1 cm.

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41 Floral initiation Free hand cross sections of the bulbs used in this study revealed that floral bud formation was similar in Hippeastrum, Habranthus and Zephyranthes bulbs (Figures 312, 3-13 and 3-14). Flower buds were formed cen trally in the apical meristem, and were initiated during the bulb’s vege tative growth but did not emer ge until the bulb reached a critical size. The elongation of flowering scap es occurred after the formation of 3 to 4 leaves on these species. We observed that flower buds were produced alternatively from each side of the meristem and formed a cro ss line in cross section (Figure 3-15). Since the apical meristem was constantly activ e (except during dormant periods) the older flower buds were located on the outer part of the bulb, identical to the leaf development pattern of these three species (Figure 3-16). The average number of leaves initiated prior to inflorescence initiation depends upon the species in Amaryllidaceae bulbs. Hartsema and Leupen (1942) found that 11 leaves were initiated before floral initiation in Amaryllis belladonna and 8 in Nerine bowdenii ; while there were 4 in Hippeastrum hybridum (Blaauw, 1931), 15 in Scadoxus multiflorus (Peters, 1971), 7 to 8 in Leucojun aestivum L. (Luyten and Van Waveren, 1938), 5 in Lycoris radiata and 10 in Lycoris squamigera (Mori and Sakanishi, 1977). We observed that there were 4 leaves formed between each inflorescence in both Habranthus and Zephyranthes bulbs (Table 3-4). In Allium cepa which is similar to Hippeastrum, Habranthus and Zephyranthes, the first stages in the development of th e leaf and inflorescence primordia in the meristematic region are similar and an inflores cence bract is indistinguishable from a leaf blade, as both are protected by an involucre and a series of bracts (Jones and Emsweller, 1936).

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42 Hippeastrum flowers are produced in groups of 2 to 6 in an umbel inflorescence and are attached to a hollow scape (around 50 cm tall) that can appear singly or more than one at the same time. A mature bulb pr oduces about 12 leaves and 3 to 4 scapes per season. Inflorescences in Habranthus and Zephyranthes possess only one flower in most cases (rarely 2 flowers) attached to a hollow scape. We observed that a mature bulb can produce up to 4 scapes at the same time (figure 3-17). Both Habranthus and Zephyranthes bulbs can flower several times during one season, their flowers last for 2 to 3 days and leaves may or not be present during flowering. Throughout this study it was observed that it takes four days for a Habranthus flower to transition from visible flower bud to an open flower (Figure 3-18). In Scadoxus bulbs, flower buds and pseudo-stems are formed side by side at the apical meristem (Figure 3-19), one of each pe r season. The inflorescence scape which is a spherical umbel (25 cm in diameter) consis ting of up to 200 flowers, emerges first and then the foliage held by a solitary pseudos tem emerges (Figure 3-20). The inflorescence lasts for 1 to 2 weeks, while the pseudo-stem lasts for a few months. Their flowering season is in late summer to early autumn. We observed that Agapanthus rhizomes did not produce inflorescence and leaves in the same region of the basal plate. Infloresce nces were formed in the apical meristem located in the central part of the basal plat e while leaves were developed from apical meristems distributed along the lateral si des of the basal plate (Figure 3-21A). Agapanthus inflorescences are umbels with 8 to 20 flowers held by stiff erect scapes that arise from the center of the rhizome (Figure 3-21B). Inflorescences last for 1 week and flowers appear during summer.

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43 Table 3-4 – Relationship between leaf and flower formati on, and number of flowers per inflorescence of plants examined Species Leaves between inflorescences Number of flowers per inflorescence Hippeastrum 4 2 to 6 Habranthus 4 1 to 2 Zephyranthes 4 1 to 2 Scadoxus 15 Up to 200 Agapanthus 8 to 20 (A) (B) Figure 3-12: Longitudinal sections of a Hippeastrum bulb. Free-hand section (A) and photomicrograph (B). Bar = 1 cm. (A) (B) Figure 3-13: Sections of Habrantus bulbs. Inner part with outer scales removed (A) and photomicrograph of a longitudinal section (B). Bar = 1 cm.

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44 (A) (B) Figure 3-14: Sections of Zephyranthes bulbs. Inner part with outer scales removed (A) and photomicrograph of a l ongitudinal section (B). Bar = 1 cm. Figure3-15: Cross section of a Habranthus bulb (stained with Safranin), showing the alignment of flower buds. Bar = 1 cm.

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45 Figure 3-16: Longitudinal section of a Habranthus bulb (stained with Safranin), showing flower buds. Bar = 1 cm. Figure 3-17 Habranthus bulb with three flower stalks. Bar = 10 cm.

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46 (A) (B) (C) (D) Figure 3-18 Development of a Habranthus flower. Flower bud (circled) emerged from soil on day 1 (A), flower stalk elongated (circled) on day 2 (B), flower stalk (circled) elongated on day 3 (C) and open flower on day 4 (D). Bar = 5 cm. (A) (B) Figure 3-19: Sections of Scadoxus bulbs. Inner part with ou ter scales removed (A) and photomicrograph of a longitudinal section (B). Bar = 1 cm.

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47 Figure 3-20: Flower stalk emerged and pseudo-stem arising from a Scadoxus bulb. Bar = 1 cm. (A) (B) Figure 3-21: Sections of Agapanthus rhizomes. Photomicrograph of an in longitudinal section (A) and flower stalk emerging from center of an Agapanthus rhizome (B). Bar = 1 cm.

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48 Meristematic region The apical meristem consists of a group of undifferentiated cells located at the apex of a shoot or a root that c ontinuously produces new cells. Th e apical meristem was first observed in 1759 by Caspar Wolff, who recogni zed it as being the center of organ and cell formation (Tooke and Battey, 2003). There are four types of meristems: single apic al cell or initial (that can be lenticular or tetrahedral), cluste r of apical cells, zoned apex and tunica corpus type. The first two types are characteristic of fern s and other lower plants, the thir d type is characteristic of gymnosperms, and the fourth type occurs in angiosperms. The five species of tropical bulbs studied have the tunica-corpus t ype of shoot meristem (Dengler, 2002). Studies of meristems reflect the technol ogy of the time period in which they were conducted. During the 19th and 20th centuries, observational analysis were made on sections of living and preserved tissue, fr om 1940 to 1970, experimental manipulations were made with microsurgery, radioisotopes, labeling and chimeric analysis. Since 1970, genetic and molecular analyses have been employed to gain information on plant meristems (Tooke and Battey, 2003). Tulip apices were described by Sass (1944) as a tunica-corpu s type, during the vegetative state and flower differentiation. The vegetative apex of th e principal axis is deeply buried in the bulb, just above the basal plate, and it is 2 to 3 mm long and 1 to 2 mm wide. The apex is a short dome 100 to 125 mm high and 300 to 375 mm broad and approximately semicircular in longit udinal section. The apical meristem of Allium cepa has also been described as a low, circ ular, dome-shaped mass of cells (Hoffman, 1933).

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49 In tulips, the tunica consists of a single la yer of cells, followed by several layers of the corpus. The leaf primordia results from di vision of cells in the first layers of the corpus and rapidly involve zone three or four cells deep. According to Rees (1972), different plants can have a tunic with more than one layer of cells. That is the case in Iris which posses a tunica of two layers of cells, and Lilium candidum which has a tunica formed by three layers of cells. This study did not identify the number of cell layers in the tunica of each species but clar ified the cellula r layout of the apical region in all five species studied. Hippeastrum, Habranthus, Zephyranthes and Scadoxus bulbs had a similar arrangement and their apical meristematic regions were located in the basal plate, as described for Allium cepa (Mason, 1979). The apical meristem and youngest leaf primordia were surrounded by scales (Figur es 3-22 to 3-29). Blaaum (1931) found that the first morphological sign of inflorescence initiation in Hippeastrum bulbs was a slight increase in the size of the apical meristem. Agapanthus rhizomes were distinctly different that Hippeastrum, Habranthus, Zephyranthes and Scadoxus as they had a centrally locate d apical meristem responsible for flower formation and several lateral meri stems located on the sides of the rhizome, which were responsible for leaf production (Figures 3-30 and 3-31).

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50 Leaf primordium Meristem Flower bud Basal plate Bulb Scale Developing Leaf Vascular channel (Swollen leaf base) Leaf primordium Developing Leaf Figure 3-22: Photomicrograph of the meristematic region of a Hippeastrum bulb. Bar = 1 cm. Leaf primordium Developing leaf Vascular channel Basal plate Meristem Leaf primordium Unmerged leaf Figure 3-23: Photomicrograph of the meristematic region of a Hippeastrum bulb. Bar =1 cm.

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51 Meristem Bulb scale Unmerged leaf Leaf primordium Basal plateSwollen leaf base Developing leaf Vascular channel Figure 3-24: Photomicrograph of the meristematic region of a Habranthus bulb. Bar = 1 cm. Leaf primordium Developing leaf Vascular channel Basal plate Meristem Leaf primordium Unmerged leaf Figure 3-25: Photomicrograph of the meristematic region of a Habranthus bulb. Bar = 1 cm.

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52 Leaf primordium Flower bud Meristem Basal plate Developing leaf Vascular channel Flower bud Figure 3-26: Photomicrograph of the meristematic region of a Zephyranthes bulb. Bar = 1 cm. Leaf primordium Basal plate Vascular channel Meristem Leaf primordium Flower bud Developing leaf Developing leaf Figure 3-27: Photomicrograph of the meristematic region of a Zephyranthes bulb. Bar = 1 cm.

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53 Developing leaf Meristem Basal plate Leaf primordium Vascular channel Leaf primordium Developing leaf Figure 3-28: Photomicrograph of the meristematic region of a Scadoxus bulb. Bar = 1 cm. Leaf primordium Leaf primordium Basal plate Vascular channel Developing leaf Meristem Developing leaf Figure 3-29: Photomicrograph of the meristematic region of a Scadoxus bulb. Bar = 1 cm.

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54 Emerged flower stalk Developing leaf Leaf primordium Basal plate Leaf primordium Leaf primordium Lateral meristem Lateral meristem Lateral meristem Leaf primordium Vascular channel Figure 3-30: Photomicrograph of the meristematic region of an Agapanthus rhizome. Bar = 1 cm. Lateral meristem Leaf primordium Basal Plate Vascular channel Developing leaf Leaf primordium Developing leaf Figure 3-31: Photomicrograph of the meristematic region of an Agapanthus rhizome. Bar = 1 cm.

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55 General anatomical and morphological arrangement Figure 3-32: Longitudinal section of a Hippeastrum bulb showing its general anatomy and morphology. Bar = 1cm. Figure 3-33: Longitudinal section of a Hippeastrum bulb showing its general anatomy and morphology. Bar = 1cm. Basal plate Flower bud Unemer g ed leaf B u l b sca l e Basal plate Bulb scale Unemerged leaf Flower bud

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56 Figure 3-34: Longitudinal section of a Zephyranthes bulb showing its general anatomy and morphology. Bar = 1cm. Figure 3-35: Longitudinal section of a Zephyranthes bulb. Bar = 1cm. Basal plate Emer g in g leaf Bulb scale Basal plate Bulb scale Unemerged leaf Flower bud

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57 (A) (B) Figure 3-36: Sections of Habranthus bulbs. Longitudinal (A) and transversal (B) sections, showing the reduced basal plat e with leaves/scales between flower buds (A) and a series of leaves/scales (B). Bar = 1cm. Figure 3-37: Longitudinal section of a Scadoxus bulb showing its general anatomy and morphology. Bar = 1cm. Bulb scale Bulb scale Pseudo stem Basal plate Basal plate Unemerged leaf Flower b ud

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58 Figure 3-38: Longitudinal section of a Scadoxus bulb showing its general anatomy and morphology. Bar = 1cm. Bulb scale Pseudo stem Basal plate Unemerged leaves

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59 Figure 3-39: Longitudinal section of an Agapanthus rhizome showing its general anatomy and morphology. Bar = 1cm. Figure 3-40: Longitudinal section of an Agapanthus rhizome showing its general anatomy and morphology. Bar = 1 cm. Unemerged leaf Emerged leaf Basal plate Emerged leaf Emerged inflorescence Unemerged leaf Rhizome Basal plate

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60 Experiment 1 The storage organs of geophytes are able to supply food reserve for rapid leaf growth at the beginning of thei r growth and reproductive cycle, after cold winters and/or dry seasons (Rees, 1972). In bulbs with hystera nthous leaves (flowers and leaves develop in separate seasons), an accumulation of storag e materials is a prerequisite for flowering (Burtt, 1970). In bulbs with synanthous leaves which are present when plants flower, the storage materials provide for early growth of photosynthetic organs and allocation of resources for flowering originate from the st orage organ and from ex isting leaves (Dafni et al., 1981). Phenological differences can be summarized into two groups based on the speed of leaf development and duration of photos ynthetic period. The rapid route is a characteristic of bulbs that produce leaves quickly in order to compensate for a short photosynthetic period or when there is a li miting factor, such as moisture (most common), light or temperature. The slow rout e is characteristic of bulbs that produce leaves slowly, under long photosynthetic period s or when moisture, light and temperature are not limiting factors (Dafni et al., 1981). Of the five species used in this morphological study, Habranthus and Zephyranthes were the least investigated by other researcher s, and since these two genera were used in the subsequent phases of this study an experi ment was designed to understand their leaf and flower production. Habranthus and Zephyranthes have perennial storage organs and synanthous leaves. According to Dafni et al. (1981) this type of bulb accumulates more reserves than required for a growth cycle which includes completi on of flowering and seed production even if the net production (by existing leaves) is insufficient. Therefore relatively slight differences are expected from year to year in the abundance of flowering.

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61 The following experiment was performed to compare leaf and flower production of Habranthus and Zephyranthes during a period of one year. Materials and Methods : On the first week of January 2005, 12 Habranthus robustus bulbs and 12 Zephyranthes bulbs were planted in 15 cm plastic pots, one bulb per pot, using a sphagnum peat based Fafard No. 2 soiless growing medium (Agawam, MA) consisting of 70% Canadian sphagnum peat, 10% perlite and 20% vermiculite. Plants were placed in a greenhouse with 11% shade, natural photoperiod and a temperature range of 31/24oC (day/night). Plants receiv ed 250ml of water and were fertilized at each irrigation with Peters Professional ‘F lorida Special’ water soluble fertilizer 20N-4.7P-16.6K (Scotts Co., Marysville, OH) with N at 150 mg.L-1. Bulbs used in this experiment were obtained from the University of Florida stock and had been grown in ground beds for a year prior the beginning of the experiment. This experiment was performed from January to December 2005. Data were collected twice a month during the entire y ear and the number of leaves and flowers present at the moment of data collection were recorded. Da ta was analyzed by taking the mean number of leaves and fl owers per total of bulbs at each observation, using the SAS statistical package version 8.02 (Cary, NC). Re sults demonstrate the performance of leaf and flower production of these two species during 2005. Results : Habranthus bulbs demonstrated a predictable performance of leaf production with more leaves at the beginning an d end of the year when not flowering and a gradual reduction of leaf emergence as flowering season occurred (Figure 3-41). Zephyranthes also produced more leaves in the be ginning and end of the year, but their flowering season did not coincide with a period of reduced leaf production. In April

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62 Zephyranthes bulbs produced the least number of l eaves followed by a gradual increase until August when a slight drop occurred and the flowering season st arted. Production of leaves increased again until October, when another reduction occurred followed by an increase of leaf emergence until the end of the year (Figure 3-42). Comparing both figures (341 and 3-42) makes it clear that these two species posses distinct patterns of leaf production. Habranthus bulbs had fewer leaves emergent when flowering, while leaves of Zephyranthes bulbs continued to emerge during flowering. Overall, Zephyranthes bulbs had a greater quantity of leaves during the entire year but with oscillations, while Habranthus bulbs demonstrated a more uniform progress with a gradual decrease and increase in leaf correlated with the flowering period. This information is valuable for the landscape use of these two species as they generate distinct appearances in a garden; Zephyranthes have copious foliage in combination with their flowers and Habranthus have few leaves when flowering.

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63 0 1 2 3 4 5 6 7 8 24681012141618202224262830323436384042444648505254 WeekNumber of Leaves / Flowers LEAVES FLOWERS Figure 3-41: Number of leav es and flowers produced by Habranthus bulbs in 2005 0 1 2 3 4 5 6 7 8 9 10 24681012141618202224262830323436384042444648505254 WeekNumber of Leaves / Flowers LEAVES FLOWERS Figure 3-42: Number of leav es and flowers produced by Zephyranthes bulbs in 2005

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64 CHAPTER 4 ENVIRONMENTAL EFFECTS – FERTIGATION FREQUENCY AND FERTILIZER RATES ON FLOWERING IN Habranthus robustus AND Zephyranthes spp The primary goal of a flower bulb grower is to produce true-to-type, essentially diseaseand insect-free plants that flow er successfully. Proper water management and fertilization are among the critical factors to accomplish that goal. Most all ornamental bulbs along with onion and garlic (related vegetable crops) are sensitive to water deficit. When the soil is ke pt relatively moist, root growth is reduced which favors bulb enlargement and consequen tly flowering, since flower production in bulbs is highly affected by bulb size in several species including Hippeastrum, Habranthus, Zephyranthes (Theron and De Hertogh, 2001) A frequent and light irrigation is commonly practi ced on both vegetable and orna mental bulbous crops, while over-irrigation is avoided since it can increase the incidence of several diseases (Hafeez, 1984). Soil water deficits inhibit leaf expansion, as nutrient uptake is reduced because of reduced transpiration rates (R ussel, 1977). In onion, transpir ation rates, photosynthesis and growth are lowered by mild water stresse s (Begun et al., 1990). Stressed onions often bulb too early, produce small-sized bulbs and ha ve an increased rate of bulb split. All these factors can reduce mark etable yields (Hegde, 1986). The soil moisture requirement of onions is influenced by seve ral factors including cultivar used, soil type and temperature, light levels and other envi ronmental factors. A crop can be grown to maturity under a soil mo isture deficit, but higher yields were

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65 associated with irrigation fre quencies that eliminated soil moisture deficits (De Lis et al., 1967). Minimizing the period when leaf and flow er tissues are wet by using proper irrigation management reduce moisture ad c onsequently the development of most fungal and bacterial caused diseases. Fertilizer form ulations also influence disease development in flower bulb crops since they can alter soil pH, therefore fe rtigation should be carefully planned in bulb production. Nutrient requirements for greenhouse a nd outdoor bulb production can be grouped into four categories according to De Hertogh and Le Nard (1993): 1) bulbs containing sufficient nutrients to produce high quality potted plants or cut flowers without additional fertilization such as Hippeastrum, Hyacinthus and Narcissus ; 2) bulbs that require either no additional fertilization or in which the application of Ca(NO3)2 can eliminate or reduce physiological disorders such as Dutch ir ises and tulips; 3) bul bs that require low fertilization programs, such as Anemone, Freesia and Liatris ; and 4) bulbs that require moderate fertilization programs, such as Dahlia, Gladiolus, Lilium and Ranunculus Fertilization programs have been evaluated on several genera of flowering bulbs to determine their effect on flowering, flower quality, bulb growth, bulb yield, and seed yield. Flowering as well as bulb growth of Iris Tulipa and Lilium were compared when plants were treated with 10.7:3.9, 12.1:5.1, 14.3: 3.9, or 17.9:3.9 meq/L of N and K. (Lee et al., 2005). Results showed that flow ering was slightly accelerated when Iris was grown with 10.7:3.9 meq/L N:K; flowering of Tulipa was promoted by 14.3:3.9 meq/L N:K; and neither growth or flowering of Lilium was significantly different between any fertilizer treatments. When 0, 30, 70, 120, 180, 250, 330, 420, and 520 kg N ha-1 were applied to

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66 Lachenalia cultivars, results demonstrated that increased rates of nitrogen had a positive influence on bulb fresh mass and circumfere nce (Engelbrecht, 2004) Three levels of fertilizer, 50, 100, and 200 mg l-1 N, were applied to Curcuma alismatifolia of the Zingiberaceae family and results showed that plants treated with the highest fertilizer level were taller, produced mo re stems per cluster, had la rger diameter rhizomes, a greater number of new rhizomes and better flower quality (Ruamrungsri et al., 2005). In onions, irrigation interval s of once a week, every 10 da ys and every other week were investigated in associat ion with fertilizer levels of 90 and 180 kg N/ha to ascertain the optimum irrigation interval and fertilizer level that pr oduced the highest yields of good quality bulbs. Results demons trated that the 10 day irri gation interval and the 90 kg N/ha rate resulted in the highest yield per ha and the greatest average bulb weight. However, no treatment interactions were de tected (Hassan, 1984). Irrigation frequencies were also investigated on Caladium tubers in a pot study and results showed that three times a week provided the best yield response compared to once a week and twice a week (Overman and Harbaugh, 1988). Extremely limited information was found in the horticultural lit erature regarding Habranthus and Zephyranthes fertigation frequencies or fe rtilizer regimes. The present study was designed to address this issue and two experiments were conducted: 1) to determine optimal fertigation frequency for these two genera in order to achieve an extended flowering season with increased number of flowers per bulb, and 2) to determine optimal fertilizer rates for bulb development. In the first experiment, different fertigati on frequencies (twice a week, once a week and every other week) were applied to Habranthus robustus and Zephyranthes spp. to

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67 determine their effect on fl owering and/or on bulb development (number of leaves, number of offsets, bulb size, bulb weight and number of bud s). In the second experiment, different fertilizer rates 0, 75, 150 and 300 ppm N, were applied to Habranthus robustus and Zephyranthes spp to verify their effect on flowering and bulb development. The Habranthus species used in this project Habranthus robustus is the most popular species in the genus; it is native to Brazil but grows very well in Florida. The Zephyranthes species used in this projects were: Z. ‘Paul Niemi’, Z. JoAnn’s Trial’, and Z. ‘Fadjar Pink’ which were hybridized by Fadjar Marta in Jakarta, Indonesia. Experiment 1 : Influence of three different fertigation treatments on number of flowers and bulb development in Habranthus robstus and Zephyranthes spp. Materials and Methods : Bulbs used in this experiment were obtained from University of Florida stock, which had been grown in ground beds for a year prior to the beginning of the experiment. On July 02, 2004, th ese bulbs were planted in 15 cm plastic pots, three bulbs per pot using sphagnum peat based Fafard No. 2 soiless growing medium (Agawam, MA) consisting of 70% Ca nadian sphagnum peat, 10% perlite and 20% vermiculite. All plants were placed in a greenhouse (Figure 41) under 11% shade, natural photoperiods and a te mperature range of 31/24oC (day/night). These plants received 250 ml of wa ter with N at 150 mg.L-1, using Peters Professional ‘Florida Special’, a water soluble fertilizer with 20N-4.7P-16.6K (Scotts Co., Marysville, OH). This experiment was conducted in two di fferent years: year 1 July 15 to December 1, 2004 (week 29 to 47), and year 2 – May 3 to December 1, 2005 (week 18 to 49). Fertigation treatments of 1) twice a week, 2) once a w eek, and 3) every other week were started on July 15, 2004 the first y ear and May 3, 2005 on the second year.

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68 There were 16 pots (48 bulbs) per treatme nt. During the growing phase of the experiment, number of flowers was recorded twice a week during both years. A flower was counted when the petals opened to expose the anthers and was removed after collection of data. At completion of each ye ar all bulbs were sectioned in order to compare bulb development and floral initia tion. Number of leaves, number of offsets produced, total fresh weight (bulb with all leaves, roots and offsets attached), fresh weight (bulb only), bulb size (diameter), and number of flower buds produced were recorded, and were subjecte d to regression analysis. Figure 4-1: Habranthus bulbs during fertigation experiment with different plastic tags in different color distinguishing the three treatments

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69 (A) (B) Figure 4-2: Habranthus bulbs being weighed after comp letion of experiment, only bulb (A) and bulb with leaves, roots and offsets (B) Results and Discussion The effects of the different fertigati on treatments on number of flowers of Habranthus robustus could not be analyzed during the first year as the data collection started after the flowerin g peak for this crop. During year 2 plants responded differently to treatments regarding the number of flowers (Figure 4-3). Plants that were fer tigated every other week flowered one week earlier than plants fertigated once and twi ce a week. This may have occurred due to drought stress. These plants ended their flow ering season two weeks earlier than plants fertigated once a week and eight weeks earlier than plants fertigated twice a week. Plants fertigated once and twice a week started flow ering during week 19 but plants fertigated twice a week flowered longer, until week 37, wh ile plants fertigated once a week stopped flowering in week 31. Peak flowering occurred much earlier in plants fertigated twice a week (week 21), than plants fertigated on ce a week (week 24) and every other week (week 26). Number of flower s produced during peak flow ering differed dramatically

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70 among the treatments, as plants fertigated tw ice a week had 41 flowers compared to 25 for plants fertigated once a week and 34 for plants fertigated every other week. Plants fertigated once and twice a week produced flowers more consistently than plants fertigated every other week, as these plants had a flush of flowers at the beginning of the season, flowered poorly during followi ng weeks, did not flower for a month and then had a second flush of flowers. Both regi mens of fertigation – watered twice or once a week seemed to be effective on Habranthus under greenhouse environment, as these regimens resulted in longer flowering pe riods and greatest number of flowers. Similar results were observed in oni ons where water deficits resulted in underdeveloped plants with reduced yields (Kadayifci et al., 2004). According to Mermoud et al. (2005) changes in the irrigation frequency si gnificantly influences the components of the water balance. A decrease in irrigation frequency causes an increase in the water storage in the root zone and a moistu re deficit in the immediate vicinity of the soil surface, which affect the crop’s performan ce. This may have occurred in this study when plants were fertigated every other week. The results obtained after comp letion of the first year de monstrated that different fertigation frequencies affected Habranthus bulb development. Regression analyses of all factors investigated (number of leaves, numb er of offsets, bulb size, total fresh bulb weight, bulb weight and number of flower buds) showed lin ear responses with coefficient of determination above 0.50. All factors incr eased as fertigation increased in year 1 (Figures 4-4 to 4-9). Number of offsets, to tal fresh bulb weight a nd bulb weight had the highest coefficient of determinations; all we re above 0.90. These results can be compared to those obtained by Overman and Harbaugh (1988) who tested different irrigation

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71 frequencies in Caladium tubers and observed that most fre quent irrigations resulted in the best yields. After completion of the second year the re sults obtained supp orted the conclusion that different fertigation syst ems affect some aspects of Habranthus bulb development as described bellow. There were no linear respon ses for number of leaves and number of flower buds (Figures 4-10 and 4-15). Number of offset s decreased as fertigation frequencies increased (r=1) (F igure 4-11). Bulb size, tota l fresh bulb weight and bulb weight increased as fertiga tion frequencies increased; how ever bulb weight (total and only bulb) had higher coefficients of determ ination; both were above 0.90 (Figures 4-12, 4-13 and 4-14). During year 1 Zephyranthes responded differently regard ing flowering to different fertigation frequencies compared to Habranthus This was evident in Zephyranthes during the first year, but not in Habranthus as both species ( Habranthus and Zephyranthes ) have different flowering seasons and Zephyranthes plants begin to flower much later than Habranthus plants (Figure 4-16). The results of year 1 showed that Zephyranthes bulbs fertigated once a week and twice a week had three gradual peaks of flow er followed by an abrupt decline. This did not happen when plants were fertigated ev ery other week. Plants fertigated once and twice a week flowered similarly which differe d from plants fertigated every other week (Figure 4-17). Plants fertigated twice a week and every other week flowered for one week more than the ones fertigated once a week. Plants fertigated twice a week had, overall, a greater

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72 quantity of flowers during the entire season, but plants fertigated every other week had more flowers during the first peak -37 fl owers in a group of 36 bulbs (Figure 4-17). During year 2 plants fertigated once a w eek and twice a week demonstrated three flowering peaks followed by an abrupt decl ine in flower production. These two groups had similar flowering patterns, which differed from plants fertigated every other week. Plants fertigated twice a week had a greater number of flowers (Figure 4-18). Plants fertigated once a week started fl owering two weeks earlier than plants fertigated twice a week and finished their flowering season one week before both plants fertigated twice a week and every other week. Plants fertigated twice a week had, overall, a greater number of flowers during the entire season, but pl ants fertigated every other week had more flowers during their first peak -39 flowers in a group of 36 bulbs. Plants that were fertigated once and tw ice a week produced their first flowers earlier in the season (week 19 for plants fertigated once a week and week 21 for fertigated twice a week), but plants under both regimens fl owered poorly for more than two months, until the first flush of flowers, on week 31. Both once and twice a week fertigation regimens seemed to be effective for flowering in Zephyrathes under greenhouse environments. Bu t fertigating twice a week would be preferable since the results showed that plants under this fertigation regimen produced more flowers. Similar experiments were performed with onion and garlic (a tunicate type and a non-tunicate type of bulb), and a once to twice a week watering regimen has also been found to be prefer able for these crops (Hanson et al., 2003; Kadayifci et al., 2004; Mermoud et al. 2005). Fert igating plants every other week reduced flowering of Zephyranthes bulbs as it resulted in a shorte r flowering season with intervals

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73 of no flowers. Similar experiments could be done in further studies using different conditions of light and temper ature in order to better de fine the ideal fertigation frequency for these two species of tropical bulbs. The treatments affected Zephyranthes bulb development in year 1. Regression analysis of all factors investigated, with the exception of number of offsets (Figure 4-20), showed linear responses with coefficient of determination above 0.50. All factors that had a linear response (number of leaves, bulb size total fresh bulb weigh, bulb weight and number of flower buds) increased as fertigati on increased during the first year (Figures 419, 4-21, 4-22, 4-23, and 4-24), similarly to the Habranthus study. Number of leaves and fresh bulb weight had highest coefficien t of determination, both were above 0.90. During year 2, number of offsets and total fresh bulb weight were the only factors that had linear responses, with coefficient of determination above 0.50 (Figures 4-25 to 430). Number of offsets decreased as fertig ation frequencies incr eased (Figure 4-26), while total fresh bulb weight increased as fertigation fr equencies increased and was highest when bulbs were fertigat ed once a week (Figure 4-28).

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74 0 0.2 0.4 0.6 0.8 1 1.2 1819202122232425262728293031323334353637 TimeNumber of flowers Fertigated twice a week Fertigated once a week Fertigated every other week Figure 4-3: Number of flowers of Habranthus robustus bulbs in year 2 as affected by fertigation frequency (n=48). Bars represent standard errors. y = 0.561x + 4.3687 R2 = 0.5711 0 2 4 6 8 01234 Fertigation frequencyNumber of leaves Mean Linear (Mean) Figure 4-4: Number of leaves of Habranthus robustus bulbs in year 1 as affected by fertigation frequency. 1=every other week 2=once a week 3=twice a week

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75 y = 2.9065x 1.204 R2 = 0.93770 2 4 6 8 01234Fertigation frequencyNUmber of offsets Mean Linear (Mean) Figure 4-5: Number of offsets of Habranthus robustus bulbs in year 1 as affected by fertigation frequency. y = 0.0845x + 3.1387 R2 = 0.85413.15 3.2 3.25 3.3 3.35 3.4 3.45 01234Fertigation frequencyBulb size Mean Linear (Mean) Figure 4-6: Bulb size of Habranthus robustus in year 1 as affected by fertigation frequency. y = 6.7565x + 32.339 R2 = 0.99190 20 40 60 01234Fertigation frequencyTotal weight Mean Linear Figure 4-7: Total weight of Habranthus robustus bulbs in year 1 as affected by fertigation frequency. 1=every other week 2=once a week 3=twice a week 1=every other week 2=once a week 3=twice a week 1=every other week 2=once a week 3=twice a week

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76 y = 1.8475x + 14.411 R2 = 0.99580 5 10 15 20 25 00.511.522.533.5Fertigation frequencyBulb weight Mean Linear (Mean) Figure 4-8: Bulb weight of Habranthus robustus in year 1 as affected by fertigation frequency. y = 0.236x + 1.3603 R2 = 0.6040 0.5 1 1.5 2 2.5 00.511.522.533.5Fertigation frequencyNumber of buds Mean Linear (Mean) Figure 4-9: Number of flower buds of Habranthus robustus bulbs in year 1 as affected by fertigation frequency. y = 0.2395x + 4.936 R2 = 0.14010 1 2 3 4 5 6 7 00.511.522.533.5Fertigation frequencyNumber of leaves Mean Linear (Mean) Figure 4-10: Number of leaves of Habranthus robustus bulbs in year 2 as affected by fertigation frequency. 1=every other week 2=once a week 3=twice a week 1=every other week 2=once a week 3=twice a week 1=every other week 2=once a week 3=twice a week

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77 y = -0.375x + 19.75 R2 = 118.4 18.6 18.8 19 19.2 19.4 19.6 00.511.522.533.5Fertigation frequencyNumber of offsets Mean Linear (Mean) Figure 4-11: Number of offsets of Habranthus robustus bulbs in year 2 as affected by fertigation frequency. y = 0.1155x + 2.8087 R2 = 0.59372.85 2.9 2.95 3 3.05 3.1 3.15 3.2 00.511.522.533.5Fertigation frequencyBulb size Mean Linear (Mean) Figure 4-12: Bulb size of Habranthus robustus in year 2 as affected by fertigation frequency. y = 6.777x + 37.928 R2 = 0.96850 10 20 30 40 50 60 70 00.511.522.533.5Fertigation frequencyTotal weight Mean Linear (Mean) Figure 4-13: Total bulb weight of Habranthus robustus in year 2 as affected by fertigation frequency. 1=every other week 2=once a week 3=twice a week 1=every other week 2=once a week 3=twice a week 1=every other week 2=once a week 3=twice a week

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78 y = 3.0715x + 8.695 R2 = 0.99730 5 10 15 20 00.511.522.533.5Fertigation frequencyBulb weight Mean Linear (Mean) Figure 4-14: Bulb weight of Habranthus robustus in year 2 as affected by fertigation frequency. y = 0.0855x + 2.7053 R2 = 0.38932.7 2.75 2.8 2.85 2.9 2.95 3 3.05 00.511.522.533.5Fertigation frequencyNumber of buds Mean Linear (Mean) Figure 4-15: Number of flower buds of Habranthus robustus bulbs in year 2 as affected by fertigation frequency. 1=every other week 2=once a week 3=twice a week 1=every other week 2=once a week 3=twice a week

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79 0 20 40 60 80 100 120 26272829303132333435363738394041424344454647 TimeTotal Number of Flowers Total flw Habrabthus Total flw Zephyranthes Figure 4-16: Total number of flowers on Habranthus and Zephyranthes bulbs from July to December 2004 (n=142) 0 0.2 0.4 0.6 0.8 1 1.2 1.4 31323334353637383940414243444546 WeekNumber of Flowers Fertigated twice a week Fertigated once a week Fertigated every other week Figure 4-17: Number of flowers of Zephyranthes spp bulbs in year 1 as affected by fertigation frequency (n=48). Bars represent standard errors.

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80 0 0.2 0.4 0.6 0.8 1 1.2 1.4 1.6 1.8 19202122232425262728293031323334353637383940414243444546WeekNumber of Flowers Fertigated twice a week Fertigated once a week Fertigated every other week Figure 4-18: Number of flowers of Zephyranthes spp bulbs in year 2 as affected by fertigation frequency (n=48). Bars represent standard errors. y = 1.0155x + 1.6587 R2 = 0.99420 1 2 3 4 5 00.511.522.533.5Fertigation frequencyNumber of leaves Mean Linear (Mean) Figure 4-19: Number of leaves of Zephyranthes spp. bulbs in year 1 as affected by fertigation frequency. 1=every other week 2=once a week 3=twice a week

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81 y = -0.125x + 9.7333 R2 = 0.12429 9.1 9.2 9.3 9.4 9.5 9.6 9.7 9.8 9.9 00.511.522.533.5Fertigation frequencyNumber of offsets Mean Linear (Mean) Figure 4-20: Number of offsets of Zephyranthes spp. bulbs in year 1 as affected by fertigation frequency. y = 0.0725x + 2.2127 R2 = 0.77072.25 2.3 2.35 2.4 2.45 2.5 00.511.522.533.5Fertigation frequencyBulb size Mean Linear (Mean) Figure 4-21: Bulb size of Zephyranthes spp. in year 1 as affected by fertigation frequency. y = 0.3455x + 32.756 R2 = 0.738132.9 33 33.1 33.2 33.3 33.4 33.5 33.6 33.7 33.8 33.9 00.511.522.533.5Fertigation frequencyTotal weight Mean Linear (Mean) Figure 4-22: Total fresh weight of Zephyranthes spp. bulbs in year 1 as affected by fertigation frequency. 1=every other week 2=once a week 3=twice a week 1=every other week 2=once a week 3=twice a week 1=every other week 2=once a week 3=twice a week

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82 y = 0.08x + 12.877 R2 = 0.979612.9 12.95 13 13.05 13.1 13.15 00.511.522.533.5Fertigation frequencyBulb weight Mean Linear (Mean) Figure 4-23: Bulb weight of Zephyranthes spp. in year 1 as affected by fertigation frequency. y = 0.196x + 2.0313 R2 = 0.62140 0.5 1 1.5 2 2.5 3 00.511.522.533.5Fertigation frequencyNumber of buds Mean Linear (Mean) Figure 4-24: Number of flower buds of Zephyranthes spp. bulbs in year 1 as affected by fertigation frequency. y = -0.0275x + 3.8513 R2 = 0.02373.65 3.7 3.75 3.8 3.85 3.9 3.95 4 4.05 01234Fertigation frequencyNumber of leaves Mean Linear (Mean) Figure 4-25: Number of leaves of Zephyranthes spp. bulbs in year 2 as affected by fertigation frequency. 1=every other week 2=once a week 3=twice a week 1=every other week 2=once a week 3=twice a week 1=every other week 2=once a week 3=twice a week

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83 y = -0.5835x + 10.111 R2 = 0.64468.2 8.4 8.6 8.8 9 9.2 9.4 9.6 9.8 10 01234Fertigation frequencyNumber of offsets Mean Linear (Mean) Figure 4-26: Number of offsets of Zephyranthes spp. bulbs in year 2 as affected by fertigation frequency. y = 0.0205x + 2.6507 R2 = 0.30582.64 2.66 2.68 2.7 2.72 2.74 01234Fertigation frequencyBulb size Mean Linear (Mean) Figure 4-27: Bulb size of Zephyranthes spp. in year 2 as affected by fertigation frequency. y = 0.85x + 34.4 R2 = 0.662834.5 35 35.5 36 36.5 37 37.5 01234Fertigation frequencyTotal weight Mean Linear (Mean) Figure 4-28: Total fresh weight of Zephyranthes spp. bulbs in year 2 as affected by fertigation frequency. 1=every other week 2=once a week 3=twice a week 1=every other week 2=once a week 3=twice a week 1=every other week 2=once a week 3=twice a week

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84 y = 0.09x + 12.657 R2 = 0.316612.65 12.7 12.75 12.8 12.85 12.9 12.95 13 13.05 01234Fertigation frequencyBulb weight Mean Linear (Mean) Figure 4-29: Bulb weight of Zephyranthes spp. in year 2 as affected by fertigation frequency. y = 0.005x + 2.55 R2 = 0.252.545 2.55 2.555 2.56 2.565 2.57 2.575 01234Fertigation frequencyNumber of buds Mean Linear (Mean) Figure 4-30: Number of flower buds of Zephyranthes spp. bulbs in year 2 as affected by fertigation frequency. Experiment 2 : Effect of different fertilizer ra tes on leaf production and bulb development in Habranthus robustus and Zephyranthes spp. Materials and Methods : On November 6, 2005, 36 Habranthus robustus and Zephyranthes spp. bulbs were selected from Univer sity of Florida stock, which had been grown in ground beds for two years prior to the beginning of this study. Each bulb had existing bulblets (offsets) removed, two leaves per Zephyranthes bulb and four leaves per Habranthus (extra leaves were also removed). A ll bulbs were weighed and the diameter measured prior to planting (Table 4-1).

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85 All bulbs were planted 15 cm plastic pots, one bulb per pot. The media used was a sphagnum peat based Fafard No. 2 soiless grow ing medium (Agawam, MA) consisting of 70% Canadian sphagnum peat, 10% perlite and 20% vermiculite. Four groups with nine pots each were placed in the same greenhouse, under 11% shade, spaced pot to pot on the bench and we re manually watered and fertilized twice a week with 250 ml of water containing 300 pp m N (0.2 oz of fertilizer per gallon of water); 150 ppm N (0.1 oz of fertilizer per gallon of water), 75 ppm N (0.05 oz of fertilizer per gallon of water) and no fertilizer (control gr oup) of the water soluble 20-1020 Peter’s Excel. A control group was included in this study to verify the performance of Habranthus and Zephyranthes bulbs under regular irrigation regimens with no fertilizer, which can occur in commercial production and/or in low maintenance landscapes. During the growing phase of the experime nt data was collected twice a month on number of leaves per bulb. Data collected after completion of the experiment included number of leaves, number of offsets produced, total fresh weight (bulb with all leaves, roots and offsets attached), fresh bulb we ight (only bulb), bulb size (diameter), and number of flower buds produced. Regression s were made to compare and contrast responses of Habranthus and Zephyranthes bulb development to the standard crop response to fertilizer rates demons trated by Janick et al. (1976). Table 4-1 Mean bulb size and weight prior to experiment Habranhus Zephyranthes Mean bulb diameter (cm) 2.6 2.5 Mean bulb weight (g) 16.56 15.85

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86 Results and Discussion Different fertilizer rates affected Habranthus bulb development. This contrasted with results obtained by Lee et al. (2005) in Lilium since neither flowering nor bulb growth was affected by different concentrations of fertilizer. Regression analysis of all f actors investigated (number of leaves, number of offsets, bulb size, total fresh bulb weight, bulb we ight and number of flower buds) showed a quadratic response with coefficients of de termination above 0.90. With the exception of number of offsets, all factors followed the standard crop respons e curve to fertilizer rates, with a gradual increase in growth followed by a luxury consumption period and a gradual decrease in growth. Bulbs fertilized with 75 and 150 ppm N ha d greater number of leaves and plants that did not receive any fertiliz er had the least number of l eaves (Figure 4-31). Number of offsets increased as fertilizer rates increas ed and was highest when bulbs were treated with 300ppm N (Figure 4-32). Bulb size and bu lb weight (both total fresh weight and bulb weight) were higher when bulbs were fe rtilized with 75 and 150 ppm N (Figures 433 to 4-35). These results slightly differ fr om those obtained by Engelbrecht (2004) in Lachenalia since largest and heaviest bulbs were obtained with higher concentrations of fertilizer. Bulbs that were fertilized with 150 ppm N produced the greatest number of flower buds, followed by those that received 75 and 300 ppm N (Figure 4-35). Different concentrations of fertilizer were tested in Iris and only the least con centrated fertilizer accelerated flowering (Lee et al., 2005). This study demonstrated that to propagate Habranthus bulbs using the separation method (removing and transplanting bulblets or offsets) the optimal fertilizer rate for

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87 mother plants would be 300 ppm of Peter’ s Excel 20-10-20, since bulbs produced more offsets under this treatment. However, to gr ow bulbs for the market optimal fertilizer rates would be 75 or 150 ppm of Peter’s Ex cel 20-10-20, which produc ed largest bulbs. Additionally, for flowering, fer tilizing plans with 150 ppm would be optimal since this treatment produced the greatest amount of flower buds. Zephyranthes bulb development was affected by the treatments. Regression analysis of all factors investigated (number of leaves, number of offsets, bulb size, total fresh bulb weight, bulb weight and number of flower buds) showed a quadratic response with coefficients of determination above 0.90, and all factors followed the standard crop response curve to fertilizer rates. Plants fertilized with 150 ppm N demonstr ated best results for number of leaves (Figure 4-37), number of offsets (Figure 4-38) and number of flower buds produced (Figure 4-42), while those that did not rece ive any fertilizer demonstrated the least favorable results. Bulb size increased as fertilizer rates increased and was largest when plants were fertilized with 300 ppm N (Figur e 4-39). This is comparable to onions that produced highest yields when fertilized with high concentrations of nitrogen (90 kg N/ha) (Hassan, 1984). Different concentrations of fertilizer were also tested in Curcuma alismatifolia and largest rhizomes were obtained with highest fertilizer concentrations (Ruamrungsri et al., 2005). For bulb weight (both total fresh weight and bulb weight) fertilizer concentrations of 75, 150, and 300 ppm N demonstrated simila r results producing the heaviest bulbs (Figures 4-40 and 4-41). These results di ffer from those obtained by Hassan (1984) in onion since heaviest bulbs were generated by highest concentrations of nitrogen.

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88 Plants fertilized with 150 ppm N pro duced the highest number of flower buds (Figure 4-42). This study demonstrated that the best fertilizer level for propagation and flowering of Zephyranthes bulbs was 150 ppm of Peter’ s Excel 20-10-20, since this treatment led to the highest numbers of of fsets and flower buds produced. However, 300 ppm of Peter’s Excel 20-10-20 was th e optimal concentration for growing Zephyranthes bulbs for the market, since it produced the largest bulbs. Both Habranthus and Zephyranthes can be classified as ei ther: bulbs that require low or moderate fertilization programs, type 3 or 4 in the nutrient requirement categories defined by De Hertogh and Le Nard (1993). y = -5E-05x2 + 0.0156x + 4.0996 R2 = 0.9117 0 1 2 3 4 5 6 0100200300400Fertilizer ratesNumber of leaves Mean Poly. (Mean) Figure 4-31: Data points, regression lines, e quations and coefficient of determination of number of leaves of Habranthus y = 2E-05x2 0.0011x + 1.7863 R2 = 0.98 0 0.5 1 1.5 2 2.5 3 3.5 0100200300400Fertilizer ratesNumber of offsets Mean Poly. (Mean) Figure 4-32: Data points, regression lines, e quations and coefficient of determination of number of offsets of Habranthus

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89 y = -2E-05x2 + 0.0075x + 2.9838 R2 = 0.8914 0 1 2 3 4 0100200300400Fertilizer ratesBulb size Mean Poly. (Mean) Figure 4-33: Data points, regression lines, e quations and coefficient of determination of bulb size of Habranthus y = -0.0007x2 + 0.2335x + 22.874 R2 = 0.96280 10 20 30 40 50 0100200300400Fertilizer ratesTotal weight Mean Poly. (Mean) Figure 4-34: Data points, regression lines, e quations and coefficient of determination of total fresh bulb weight of Habranthus y = -0.0006x2 + 0.1893x + 17.137 R2 = 0.91330 10 20 30 40 0100200300400Fertilizer ratesBulb weight Mean Poly. (Mean) Figure 4-35: Data points, regression lines, e quations and coefficient of determination of fresh bulb weight of Habranthus

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90 y = -3E-05x2 + 0.0112x + 1.6904 R2 = 0.94120 1 2 3 4 0100200300400Fertilizer ratesNumber of buds Mean Poly. (Mean) Figure 4-36: Data points, regression lines, e quations and coefficient of determination of number of flower buds of Habranthus y = -5E-05x2 + 0.0169x + 4.2895 R2 = 0.80480 1 2 3 4 5 6 7 0100200300400Fertilizer ratesNumber of leaves Mean Poly. (Mean) Figure 4-37: Data points, regression lines, e quations and coefficient of determination of number of leaves of Zephyranthes y = -4E-05x2 + 0.0162x + 0.8618 R2 = 0.99330 0.5 1 1.5 2 2.5 3 0100200300400Fertilizer ratesNumber of offsets Mean Poly. (Mean) Figure 4-38: Data points, regression lines, e quations and coefficient of determination of number of offsets of Zephyranthes

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91 y = -2E-06x2 + 0.0012x + 2.556 R2 = 12.5 2.55 2.6 2.65 2.7 2.75 2.8 0100200300400Fertilizer ratesBulb size Mean Poly. (Mean) Figure 4-39: Data points, regression lines, e quations and coefficient of determination of bulb size of Zephyranthes y = -0.0001x2 + 0.0435x + 18.231 R2 = 0.9050 5 10 15 20 25 0100200300400Fertilizer ratesTotal weight Mean Poly. (Mean) Figure 4-40: Data points, regression lines, e quations and coefficient of determination of total fresh bulb weight of Zephyranthes y = -2E-05x2 + 0.0093x + 11.593 R2 = 0.87811.4 11.6 11.8 12 12.2 12.4 12.6 12.8 0100200300400Fertilizer ratesBulb weight Mean Poly. (Mean) Figure 4-41: Data points, regression lines, e quations and coefficient of determination of fresh bulb weight of Zephyranthes

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92 y = -1E-05x2 + 0.0057x + 2.8557 R2 = 0.91540 1 2 3 4 0100200300400Fertilizer ratesNumber of buds Mean Poly. (Mean) Figure 4-42: Data points, regression lines, e quations and coefficient of determination of fresh bulb weight of Zephyranthes

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93 CHAPTER 5 ENVIRONMENTAL EFFECTS – LIGHT LE VELS ON FLOWERING AND BULB DEVELOPMENT IN Habranthus robustus AND Zephyranthes SPP. Light intensity and photoperiod influen ce several physiological processes of flowering bulbs, including photosynthesis, flower quality, flower abortion and abscission (De Hertogh and Le Nard, 1993). Photosynthesis is a vital process in which plants utilize sun light and carbon dioxide in the air to pr oduce sugars by converting light energy into chemical energy. Light intensity and quality al so directly affects se veral important plant quality characteristics, such as number of flowers produced, foliage color and overall plant appearance. In many cases, effects can be enhanced by selecting optimal temperature, nutrient and soil moisture levels (Gest, 2002). Photoperiodism was first de scribed by Garner and Allard (1920, 1923) in a study that classified plants as e ither long day or short day plan ts for flowering. Plants that required more than 12 hours of light pe r 24-hour cycle were considered long day (LD)/short night plants, and plan ts that required more than 12 hours of dark per 24-hour cycle were considered short day (SD)/long night plants. Onions ( Allium cepa) are considered LD plants (Lercari, 1982). Their bulb formation and subsequent growth are influenced by photoperiod (Scully et al., 1945) and temperature (B rewster, 1977), with inflorescence initiation accelerated under long photoperiods (Brewster, 1992). Bulb growth and development in garlic is al so largely dependent on photoperiod and temperature prevailing during the gr owth period (Ledesma et al., 1980).

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94 Habranthus and Zephyranthes do not require a critical amount of light or dark period in order to flower. These plants are not qualitative since they do not flower in response to short or long days Basic light requirements for fl ower bulbs in the landscape have been broadly defined as full sun, part ial shade or full shade, and according to Haynes et al. (2001), both Habranthus and Zephyranthes are considered full sun plants. Alstroemeria rhizomes were grown under shade levels of 0, 50%, and 70% and shading increased flower stem length but ha d no effect on the number of flowers per plant (Zizzo et al. 1992). However, when Oxalis braziliensis was grown under the shade levels of 35%, 55%, and 75%, number of flower s increased as shade increased (Suh et al. 2003). Similar experiment conducted using banana rhizomes grown under 0, 30%, 60% and 80% shade revealed that only 80% shade affected rhizome development (Israeli et al., 1995). This investigation was conducted to determine if Habranrhus and Zephyranthes were full sun plants and identify optimu m light levels for flowering and bulb development of these two species during commercial production. The Habranthus species used in this study was Habranthus robustus, and three Zephyranthes cultivars evaluated were ‘Paul Niemi’, ‘J oAnn’s Trial’, and ‘Fadjar Pink’. Materials and Methods Experiment 1 : Effect of light levels on fl owering and bulb development of Habranthus robustus To determine optimal light levels and verify their effect on flowering and bulb development (size, weight, number of buds), plants were grown in full sun, 30% shade and 60% shade. Bulbs used in this experiment were obtained from the University of Florida stock, where they had been grown in ground beds for a year prior to the beginning of the experiment. On July 4, 2004, bulbs were planted in 30 cm

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95 plastic pots, three bulbs per pot using sphagnum peat based Fafard No. 2 soiless growing medium (Agawam, MA) consisting of 70% Ca nadian sphagnum peat, 10% perlite and 20% vermiculite. Plants were placed in th ree different locations under different amounts of light: ambient outdoor – full sun, and two different sh adehouses (covered by black shade cloth) with 30% and 60% shade. Th ere were 16 pots (48 bulbs) per treatment. Photometric readings and temperatures of th ese three locations are listed in table 5-1. Plants were fertigated with 500ml of water containing N at 150 mg.L-1 using Peters Professional ‘Florida Special’ water solubl e fertilizer 20N-4.7P-16.6K, from Scotts Co., Marysville, OH twice a week. However, since plants were located in open environments rainfall was also monitored. A summary of mont hly rainfall totals in the Gainesville area for 2004 and 2005 is listed in table 5-2. This experiment was conducted in two differe nt years: year 1: July 15 to December 12 2004, from week 29 to 47, and year 2: May 3 to December 13 2005, from week 18 to 49. Light level treatments in both years were 1) full sun, 2) 30% shade, and 3) 60% shade. At completion of each yearly study all bulbs were sectioned in order to compare bulb development and floral initiation be tween treatments. Data collected after completion of the experiment included numbe r of leaves, number of offsets produced, total fresh weight (bulb with all leaves, roots and offsets attached), fresh bulb weight (only bulb), bulb size (diamete r), and number of flower buds produced, and regression analysis were made to compare Habranthus and Zephyranthes growth to the standard plant growth curve demonstrat ed by Janick et al. (1976).

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96 Experiment 2 Effect of light levels on fl owering and bulb development of Zephyranthes spp. To determine optimal light levels and verify their effect on flowering and bulb development (size, weight, number of buds), plants were grown under full sun, 30% shade and 60% shade. Three types of bulbs were used in this present experiment: Zephyranthes ‘Paul Niemi’, Z ‘JoAnn’s Trial’, and Z ‘Fadjar Pink’. Bulbs were obtained from the University of Florida stoc k, were they had been grown in ground beds for a year prior to the beginning of the experiment. This experiment was conducted in the sa me manner as experiment 1, using the same type of pots, media, fertilizer and placing plants in the same shadehouses. The experiment was initiated and terminated on same dates; data were collected and analyzed as experiment 1. Table 5-1 Photometric readings and temper atures at three locations / treatments Treat. Location Light Level (Mols.m2.sec) Average sunny day at noon (in September) Temperature oC Averagebetween three readings Temperature -oC Averagebetween three readings 1 Full sun 1500 31.67 28.33 2 30% Shade 1050 30.82 27.94 3 60% Shade 600 30.02 27.01 (A) (B) (C) Figure 5-1: Plants under three different treatments: full sun (A), 30% shade (B) and 60% shade (C)

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97 Table 5-2 Rainfall monthly summary in in ches for Gainesville area in 2004 and 2005 according to the Florida Automated Weather Network Jan Feb Mar Apr MayJun Jul Aug Sep Oct Nov Dec 2004 1.33 5.65 1 1.58 0.64 6.86 10.035.94 12.94 1.3 2.65 1.87 2005 1.98 2.16 5.62 4.14 4.52 7.9 5.43 4.66 0* 6.54 4.02 22.3* *these values may reflect data inaccuracies by FAWN. Results and Discussion Experiment 1 : The effects of the different lig ht level treatments on number of flowers of Habranthus robustus could not be analyzed during the first year as the data collection started af ter the flowering peak for this crop. During year 2, data collected showed that treatments did affect flower number. Plants in 30% shade had more open flowers dur ing the entire flowering season than in the other light levels. These results differ from those obtained by Zizzo et al. (1992) who observed that shading increased flower stem length but had no effect on the number of flowers per plant in Alstroemeria bulbs. Results also differ from those observed by Suh et al. (2003), who reported that number of Oxalis braziliensis flowers increased as shade levels increased. The three groups of plants started floweri ng during week 21, however, plants in full sun and in 30% shade ceased flowering after week 32, while the ones in 60% shade ceased flowering after week 27. These results also differ from thos e obtained by Zizzo et al. (1992) with Alstroemeria since they demonstrated that plants under higher level of shading had an extended flowering season. Flowering of plants in full sun peaked duri ng week 23, the same w eek that plants in 30% shade produced their first of two peaks; however, plants in full sun had less than

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98 50% of the flowers produced by plants in 30% shade. Furthermore, plants in 30% shade had a second flowering peak during week 27 w ith a great number of flowers (20% higher than the first peak of the full sun group). Plants in 60% shade also had two peaks of flowering; however, their flowering season wa s much shorter than plants in the other shade levels (Figure 5-2). The results obtained after comp letion of the first year de monstrated that different light levels affected Habranthus bulb development. Regressi on analysis of all factors investigated (number of leaves, number of offs ets, bulb size, total fresh bulb weight, bulb weight and number of flower buds) showed linear responses w ith coefficient of determination above 0.89. All factors, with th e exception of number of leaves, increased as light levels increased during year 1 (Figures 5-3 to 5-8). Number of leaves decreased as light levels increased (F igure 5-3). This may have occurred because plants in full sun received more solar radiation and fewer leaves were needed to produce sufficient energy via phot osynthesis; while plants in 60% shade received less solar radiat ion and responded by producing more leaves for photosynthetic production. These results differ from those obt ained by Sush et al. (2003) who tested different shade levels (35-55-75%) in Oxalis braziliensis and observed that number of leaves was increased by shade levels as comp ared to natural light. However, our results can be compared to those obtained by Israe li et al. (1994) who tested the effect of different shade levels (0-30-60-80%) on grow th of banana and observed that shading reduced the rate of leaf emergence. Results obtained on bulb development during the first year contrasted with those obtained by Rahim and Fordha m (1990) who tested the effect of shade on the

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99 development of garlic bulbs and observed th at bulbs grown under 50% shade were more developed (regarding number of cloves fo rmed) than those grown under 60 and 75% shade. Our results also contrasted with those obtained by Barkham (1980) who observed that Narcissus bulbs were heavier under the highest levels of canopy shade. After completion of the first year it was observed that the root system of plants grown under the different light levels diffe red. Plants grown under full sun had shorter roots with no ramifications; plants grown under 30% shade had medium roots with few ramifications, and plants grown under 60% shade had very long roots with several ramifications (Figure 5-9). This could have b een due to higher soil temperature in full sun pots. The results obtained after completion of the second year emphasized results obtained during the first year that different light levels affect Habranthus bulb development. Regression analysis of all f actors investigated (number of leaves, number of offsets, bulb size, total fresh bulb wei ght, bulb weight and nu mber of flower buds) showed linear responses with coefficien t of determination above 0.80. All factors increased as light levels incr eased during year 1 (Figures 510 to 5-15).Number of leaves increased as light levels in creased during the second year which did not happen during year 1 (Figure 5-10). Comparing the slopes of the li near responses it could be observed that the increase in number of offsets and number of flower buds was more accentuated in the second year; total fresh bulb weight increase was mo re accentuated during the first year; and bulb size and fresh bulb we ight increase were similar during both years.

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100 Experiment 2 : Effect of light levels on fl owering and bulb development of Zephyranthes spp. Light levels affected the number of Zephyranthes flowers for this species during the first year This result differed from Habranthus as Zephyranthes plants begin to flower much later than Habranthus plants (Figure 4-16) and in this case, data collection started prior to the flowering peak for this crop. Plants grown under all three treatments de monstrated gradual peaks of flowering followed by an abrupt decline duri ng week 33 (Figures 5-16 and 5-17). In the first year plants grown under 30% shade demonstrated a more uniform flowering season compared to plants in the other two light levels, with a more gradual increase in flowers leading to a peak and only one week with no flowers. Additionally, these plants had the highest number of flower s in almost all weeks. Plants grown under shade (both 30% and 60%) flowered until w eek 42, while plants grown under full sun flowered until week 44, two weeks more than the other two groups. Plants grown under full sun and 60% shade also demonstrated gra dual peaks of flowering (similar to the ones under 30% shade), however here, these peaks were followed by gaps of one or two weeks with no flowers, whereas plants under 30% shade produced flowers continually (Figure 5-16). In the second year, plants grown unde r 30% shade again had a more uniform flowering season compared to the other two groups. However, during year 2 the number of flowers during the first two peaks (week 32 and 37) were very similar among all treatments, differing from the previous seas on. During the third peak of flowering (week 41) plants under full sun had the highest num ber of flowers. Plants grown under full sun and 30% shade started flowering during week 24; both demonstrated a gap of two weeks

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101 without flowers, and flowered again on week 27, when 60% shade plants begun to flower. Plants grown under full sun flowered until week 44, two weeks more than plants under any shade level. Plants grown under the three treatme nts presented gaps without flowering between peaks and the beginning of the next gradual incr ease in number of flowers, but it was more accentuated on pl ants under full sun and under 60% shade (Figure 5-17). Results obtained during year 1 were very similar to year 2. These results differ from those obtained by Cavins and Dole (2002) who tested different shade levels (0-3060%) in Narcissus and Tulipa and observed that increasing shade increased stem length but did not influenced flowering percentage. Regarding bulb development, treatments affected Zephyranthes bulb development during both years. Regression analysis of a ll factors investigated (number of leaves, number of offsets, bulb size, total fresh bul b weight, bulb weight and number of flower buds) showed linear responses with coeffici ent of determination above 0.89. All factors increased as light levels incr eased during both year 1 and year 2 (Figures 5-18 to 5-23). Number of leaves increased during both year 1 and 2; however, it was more accentuated during the first year when compari ng the slope of the linear responses. These results differ from those obtained by Sush et al. (2003) who tested di fferent shade levels (35-55-75%) in Oxalis braziliensis and observed that number of leaves was increased by shade levels as compared to full sun. Number of offsets, bulb si ze and total fresh bulb weight increased similarly during year 1 and 2. Bulb weight increase was mo re accentuated during the first year, while

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102 number of flower buds increase was more accentuated during the second year, when comparing the slopes of the linear responses. After completion of the experiment it coul d be observed that the root system of these plants were distinct under the diffe rent treatments and was very similar to observations of Habranthus bulbs. Plants grown under full sun had shorter roots with no ramifications, plants grown under 30% shade had medium roots with few ramifications, while plants grown under 60% shade had very long roots with several ramifications. 0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 2021222324252627282930313233WeekNumber of Flowers Full Sun 30% Shade 60% Shade Figure 5-2: Number of flowers of Habranthus robustus bulbs in year 2 as affected by light levels (n=48). Bars represent standard errors.

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103 y = -2.5342x + 4.48 R2 = 0.985 0 1 2 3 4 0%50%100%150% Light levelsNumber of leaves Mean Linear (Mean) Figure 5-3: Number of leaves of Habranthus robustus bulbs in year 1 as affected by light levels. y = 7.8378x + 6.5667 R2 = 0.89920 5 10 15 20 0%50%100%150%Light levelsNumber of offsets Mean Linear (Mean) Figure 5-4: Number of offsets of Habranthus robustus bulbs in year 1 as affected by light levels. y = 0.3297x + 2.8358 R2 = 0.9308 2.9 3 3.1 3.2 0%50%100%150% Light levelsBulb size Mean Linear (Mean) Figure 5-5: Bulb size of Habranthus robustus in year 1 as aff ected by light levels.

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104 y = 11.642x + 40.467 R2 = 0.992 40 45 50 55 0%50%100%150% Light levelsTotal weight Mean Linear (Mean) Figure 5-6: Total weight of leaves of Habranthus robustus bulbs in year 1 as affected by light levels. y = 7.0872x + 11.861 R2 = 0.9969 0 5 10 15 20 0%50%100%150% Light levelsBulb weight Mean Linear (Mean) Figure 5-7: Bulb weight of Habranthus robustus in year 1 as affected by light levels. y = 0.9284x + 1.554 R2 = 0.9455 0 1 2 3 0%50%100%150% Light levelsNumberof buds Mean Linear (Mean) Figure 5-8: Number of flower buds of Habranthus robustus bulbs in year 1 as affected by light levels.

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105 Figure 5-9: Root condition of Habranthus bulbs, under different treatments, after completion of experiment 1. From left to right full sun, 30% shade and 60% shade y = 1.0357x + 3.2604 R2 = 0.99750 1 2 3 4 5 0%20%40%60%80%100%120%Light levelsNumber of leaves Mean Linear (Mean) Figure 5-10: Number of leaves of Habranthus robustus bulbs in year 2 as affected by light levels. y = 16.78x + 5.8733 R2 = 0.9217 0 5 10 15 20 25 0%50%100%150% Light levelsNumber of offsets Mean Linear (Mean) Figure 5-11: Number of offsets of Habranthus robustus bulbs in year 2 as affected by light levels.

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106 y = 0.5534x + 2.8975 R2 = 0.9463 3.1 3.2 3.3 3.4 3.5 0%20%40%60%80%100%120%Light levelsBulb size Mean Linear (Mean) Figure 5-12: Bulb size of Habranthus robustus in year 2 as aff ected by light levels. y = 13.852x + 44.273 R2 = 0.8428 0 20 40 60 80 0%50%100%150% Light levelsTotal weight Mean Linear (Mean) Figure 5-13: Total weight of Habranthus robustus bulbs in year 2 as affected by light levels. y = 7.8191x + 10.986 R2 = 0.8309 0 5 10 15 20 0%50%100%150% Light levelsBulb weight Mean Linear (Mean) Figure 5-14: Bulb weight of Habranthus robustus in year 2 as affected by light levels.

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107 y = 0.4145x + 2.1412 R2 = 0.97442.2 2.25 2.3 2.35 2.4 2.45 2.5 2.55 2.6 0%20%40%60%80%100%120%Light levelsNumber of buds Mean Linear (Mean) Figure 5-15: Number of flower buds of Habranthus robustus bulbs in year 2 as affected by light levels. 0 0.2 0.4 0.6 0.8 1 1.2 242526272829303132333435363738394041424344 TimeNumber of Flower Full Sun 30% Shade 60% Shade Figure 5-16: Number of flowers of Zephyranthes spp bulbs in year 1 as affected by light levels (n=48). Bars represent standard errors.

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108 0 0.2 0.4 0.6 0.8 1 1.2 242526272829303132333435363738394041424344 WeekNumber of Flower Full Sun 30% Shade 60% Shade Figure 5-17: Number of flowers of Zephyranthes spp bulbs in year 2 as affected by light levels (n=48). Bars represent standard errors. y = 0.4876x + 3.4875 R2 = 0.92013.6 3.7 3.8 3.9 4 4.1 0%50%100%150%Light levelsNumber of leaves Mean Linear (Mean) Figure 5-18: Number of leaves of Zephyranthes spp bulbs in year 1 as affected by light levels. y = 1.9604x + 7.7027 R2 = 0.89738 8.5 9 9.5 10 0%50%100%150%Light levelsNumber od offsets Mean Linear (Mean) Figure 5-19: Number of offsets of Zephyranthes spp bulbs in year 1 as affected by light levels.

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109 y = 0.1114x + 2.6218 R2 = 0.99992.64 2.66 2.68 2.7 2.72 2.74 0%50%100%150%Light levelsBulb size Mean Linear (Mean) Figure 5-20: Bulb size of Zephyranthes spp in year 1 as affected by light levels. y = 4.1319x + 47.154 R2 = 0.954148 49 50 51 52 0%50%100%150%Light levelsTotal weight Mean Linear (Mean) Figure 5-21: Total weight of Zephyranthes spp bulbs in year 1 as a ffected by light levels. y = 0.4955x + 12.532 R2 = 0.999112.6 12.7 12.8 12.9 13 13.1 0%50%100%150%Light levelsBulb weight Mean Linear (Mean) Figure 5-22: Bulb weight of Zephyranthes spp in year 1 as aff ected by light levels.

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110 y = 0.0781x + 2.9229 R2 = 0.99492.94 2.96 2.98 3 3.02 0%50%100%150%Light levelsNumber of buds Mean Linear (Mean) Figure 5-23: Number of flower buds of Zephyranthes spp bulbs in year 1 as affected by light levels. y = 4.9355x + 1.3368 R2 = 0.99850 1 2 3 4 5 6 7 0%20%40%60%80%100%120%Light levelsNumber of leaves Mean Linear (Mean) Figure 5-24: Number of leaves of Zephyranthes spp bulbs in year 2 as affected by light levels. y = 16.948x 1.79 R2 = 0.94980 5 10 15 20 0%50%100%150%Light levelsNumber of offsets Mean Linear (Mean) Figure 5-25: Number of offsets of Zephyranthes spp bulbs in year 2 as affected by light levels.

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111 y = 0.3865x + 2.4469 R2 = 0.94352.5 2.55 2.6 2.65 2.7 2.75 2.8 2.85 0%50%100%150%Light levelsBulb size Mean Linear (Mean) Figure 5-26: Bulb size of Zephyranthes spp in year 2 as affected by light levels. y = 41.254x + 23.643 R2 = 0.99990 10 20 30 40 50 60 70 0%50%100%150%Light levelsTotal weight Mean Linear (Mean) Figure 5-27: Total weight of Zephyranthes spp bulbs in year 2 as a ffected by light levels. y = 3.6149x + 10.564 R2 = 0.91990 5 10 15 0%50%100%150%Light levelsBulb weight Mean Linear (Mean) Figure 5-28: Bulb weight of Zephyranthes spp in year 2 as aff ected by light levels.

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112 y = 0.4511x + 2.6866 R2 = 0.87692.8 2.85 2.9 2.95 3 3.05 3.1 3.15 3.2 0%20%40%60%80%100%120%Light levelsNumber of buds Mean Linear (Mean) Figure 5-29: Number of flower buds of Zephyranthes spp bulbs in year 2 as affected by light levels.

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113 CHAPTER 6 CONCLUSIONS When talking about bulbs, the first ones that come to our minds are members of the genera Tulipa Hyacinthus, Narcissus and other temperate zone favorites. However, there is a large selection of tropical and subtr opical bulbous plants th at produce great spring and summer color, are easy to grow, and are suitable for Florida's climate; some examples include genera used in this study: Hippeastrum, Habranthus, Zephyranthes, Scadoxus and Agapanthus. The cultivation of ornamental bulbous crops is no longer limited to countries with a moderate climate. The production of high-qua lity flower bulbs in warm regions has become important during the last decades. Ho wever, the tropical and subtropical bulb market is practically non-exis tent in Florida, because most species are very slow to produce, some have viral problems, many cons umers are not sure how to grow them, and research is needed for this branch of the ornamental industry. This study was designed to address some of these issues, raise both homeowners and industry awareness of this group of plants; provide growers with impr oved methodology and guidelines to facilitate the production and commercialization of two genera of tropical bulbs: Habranthus and Zephyranthes The morphology of five speci es of tropical geophytes Hippeastrum hybridum, Scadoxus multiflorus Agapanthus africanus Habranthus robustus and Zephyranthes spp were compared and contrasted in chapter 3. Hippeastrum Habranthus and Zephyranthes were true tunicate bulbs with similar scales thickness, similar shapes and a brown papery

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114 tunica covering the bulb. Hippeastrum bulbs had the largest diameter and height, and Habranthus were slightly larger than Zephyranthes bulbs. Their leaves were simple, linear, with entire margin s and parallel venation, and emerged alternately (leaves originated on either side of the meristem) from the center of the bulb. Flower buds were formed alternately on each side of the apic al meristem (forming a cross line in cross section), and four leaves we re formed between each infl orescence/flower initiation. Scadoxus was a true tunicate bulb covered by a brown papery tunica with a thick rhizomatous structure at its base. Its size was comparable to Hippeastrum bulbs; however, the scales were thicker than those from Hippeastrum Habranthus and Zephyranthes Leaves had distinct ob long shapes compared to Hippeastrum Habranthus and Zephyranthes and arose in a cluster from the center forming a pseudostem (a false stem formed by the sheathing and overlappi ng of the leaf basis). Flower buds were formed centrally in the apical meristem, a nd an inflorescence stalk emerged first followed by the leaf pseudo-stem. Agapanthus was a rhizome with simple, linear leaves with entire margins and parallel venation, similar to those of Hippeastrum Habranthus and Zephyranthes, that arose alternatively from lateral meristems (leav es form on either side of the meristem). Flower buds were formed centra lly in the apical meristem. Hippeastrum Scadoxus Habranthus and Zephyranthes had similar meristematic arrangements and apical meristems were responsible for both leaf and flower initiation; in Agapanthus africanus flowers were formed at apical meristems and leaves arose from lateral meristems. Of the five species examin ed in the morphological study, Habranthus and Zephyranthes were the least investigated by other re searchers, and since they were used

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115 in the subsequent phases of this study an experiment was designed to understand their leaf and flower production. It was observed that these two species had distinct patterns of leaf production. Habranthus bulbs had fewer leaves emerge when flowering, while leaves of Zephyranthes bulbs continue to emerge during flowering. Overall, Zephyranthes bulbs had a greater number of leaves during the entire year but with oscillations, while Habranthus bulbs demonstrated a more uniform progress with a gradual decrease and increase in leaf production corre lated with flowering periods. In chapter 4 two experiments compared the responses of Habranthus robustus and Zephyranthes spp. to fertigation frequencies and fe rtilizer rates. Experiment 1 tested fertigation frequencies of twice a week, once a week and every other week on both species. Fertigation regimens of once or tw ice a week seemed to be effective for flowering of Habranthus robustus however, when fertigated every other week they had a shorter flowering period which begun and finished earlier. Plan ts fertigated twice a week had a flowering peak sooner and 40% more flowers compared to the other two groups. Regarding Habranthus bulb development, regression analysis of all factors investigated (number of leaves, number of offs ets, bulb size, total fresh bulb weight, bulb weight and number of flower buds) showed lin ear responses and all factors increased as fertigation frequencies increased during year 1. During year 2, similar responses were obtained for number of leaves, bulb size, total fresh bulb weight and bulb weight; however, there were no linea r responses for number of offsets and flower buds. Fertigation regimens of once and twice a w eek were also effective for flowering of Zephyranthes during both year 1 and 2 under greenhouse environments. However,

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116 fertigating twice a week would be preferable since the result s showed that plants under this fertigation regimen produced more flowers. Regarding Zephyranthes bulb development, regressi on analysis of all factors investigated, with the exception of number of offsets, showed linear responses and increased as fertigation frequencies increased during the first year, similarly to the Habranthus study. During year 2, number of offset s and total fresh bulb weight were the only factors that had linear responses. Experiment 2 tested bulb development responses of Habranthus and Zephyranthes when plants were fertilized with 20-1020 at rates of 0, 75, 15 0, 300 ppm N. With the exception of number of offsets, all factors investigated in Habranthus bulbs had a gradual increase in growth followed by a luxury c onsumption period and a gradual decrease in growth; following the standard crop response curve to fertilizer ra tes demonstrated by Janick et al. (1976). Bulbs fertilized with 75 and 150 pp m N had greater number of leaves; number of offsets increased as ferti lizer rates increased and was highest when bulbs were treated with 300ppm N; bulb size and bulb weight (both total fresh weight and bulb weight) were higher when bulbs were fertilized with 75 and 150 ppm N. These results demonstrated that optimum fertilizer rates for Habranthus bulb propagation by separation was 300 ppm of N using Peter’s Excel 20-10-20, since bulbs produced more offsets under this treatment. However, the op timum fertilizer rates for growing bulbs to the market were 75 or 150ppm of Peter’s Excel 20-10-20, which produced largest bulbs. Additionally, optimal fertilizer rates for fl owering would be 150ppm since this treatment produced the greatest amount of flower buds.

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117 In chapter 5 an experiment compared the responses of Habranthus robustus and Zephyranthes hybrids to light levels of full sun, 30% and 60% shade. Results demonstrated that both Habranthus and Zephyranthes flowered for a longer period under full sun and 30% shade but both species had more flowers when grown under 30% shade. These results contrasted to the responses of Alstroemeria bulbs as their flower stem length increased when shading was increased but the number of flowers per plant did not (Zizzo et al., 1992). Regarding Habranthus bulb development, all factor s investigated (number of leaves, number of offsets, bulb size, total fr esh bulb weight, bulb weight and number of flower buds) showed linear responses and increas ed as light levels increased during both year 1 and year 2. The only exception was nu mber of leaves which decreased as light levels increased during the first year. Zephyranthes bulb development was also affected by the treatments. All factors investigated s howed linear responses and increased as light levels increased during both year 1 and year 2. The root system of both Habranthus and Zephyranthes were affected by the treatments. Plants grown under full sun had s horter roots with no ramifications, plants grown under 30% shade had medium roots with few ramifications, while plants grown under 60% shade had very long root s with several ramifications. From this study it was concluded that pr eferred conditions for both species were fertigation twice a week with a fertilizer rate of 150 ppm N. and grown in full sun. However, further research with Habranthus and Zephyranthes should be considered. Fertigation frequency treatments could be applied to these plants under different conditions of light and temperat ure from those used in this st udy. Furthermore, fertilizer

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118 with different concentrations of nitrogen, phosphorus and potassium could be used. The current research on these two species is somewhat limited and would benefit from additional investigations.

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119 LIST OF REFERENCES Albrechtov J., Dueggelin M., Duerrenberg er M. and Wagner E. 2004. Changes in the geometry of the apical meristem and c oncomitant changes in cell wall properties during photoperiodic induction of flowering in Chenopodium rubrum New Physiol., 163(2): 263. Barkham, J.P. 1980. Population dyna mics of the wild daffodil ( Narcissus pseudonarcissus ). J. Ecol., 68: 635-664. Begun, R.A., Mallik, S.A, Rahman, M., A nowar, M.N. and Khan, M.S. 1990. Yield response of onion as influenced by different soil moisture regimens. Bangladesh J. Agr. Res., 15: 64-69. Bernier, G. 1988. The control of floral ev ocation and morphogenesis. Annu. Rev. Plant Physiol. Plant Mol. Biol. 39: 175-219. Blaaum, A.H. 1931. Orgaanvorming en periodiciteit van Hippeastrum hibridum Akademie van Wetenschappen Amsterdan, 29: 1-90 (with English Summary). Black, R.J., Bussey, N.C. and Burch D. 1990. Bulbs for Florida. Florida Cooperative Extension Service, Institute of Food a nd Agricultural Sciences, University of Florida, circular 522. Gainesville, Florida. Brewster, J.L. 1977. The physiology of the onion. Hort. Abst., 47: 17-23. Brewster, J.L. 1982. Effects of photoperiod, n itrogen and nutrition and temperature on inflorescence initiation and development in onion ( Allium cepa ). Ann. Bot., 51(4): 429-440. Bryan, J.E. 1989. Bulbs, volume I, A-H. Timber Press. Portland, Oregon. Burtt, B.L. 1970. The evolution and taxonomic significance of subterranean ovary in certain Monocotyledons. Isr. J. Bot., 19: 77-90. Caldwell, J and Wallace, T.J. 1955. Biological fl ora of the British Isles. J. Ecol, 43 (1): 331-341. Cavins, T.J. and Dole, J.M. 2002. Precooling, pl anting depth, and shade affect cut flower quality and perennialization of field-grown spring bulbs. HortScience, 37 (1): 7983.

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127 BIOGRAPHICAL SKETCH Camila do Amaral Brito was born on Nove mber 26, 1974, in So Paulo, Brazil, to Jos Fernando and Snia Brito. Camila gra duated from Colgio Objetivo (High School) in 1992 and two years later started her undergra duate degree in architecture and urbanism at the University of So Paulo, Brazil. Wh ile pursuing her degree, she has spent eight months in Vancouver, Canada, and visited se veral countries around the world. On her last year at the University, Camila worked as an intern in a landscap e design company and became fascinated by the plasticity of plan ts. Her experience on this company, under the supervision of the company’s owner, changed her life and the focus of her career. After earning her bachelor’s degree from the Univ ersity of So Paulo, she specialized in landscape architecture and worked for three years on this field. Camila was married to Luiz Augusto de Castro e Paula in Janua ry 2003 and changed her name to Camila do Amaral Brito de Castro e Paula. She and he r husband moved to Gainesville, Florida, in May of the same year and both became gradua te students at the Univ ersity of Florida. Camila was interested in environmental hort iculture and pursued her graduate degree under the guidance of Dr. Rick Schoellhorn; a year later Dr. Dennis McConnell became her major professor, while her husband pursued his graduate degree in animal sciences under Dr. Peter Hansen. Camila then earne d her Master of Sc ience in May of 2006.


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MORPHOLOGICAL ANALYSIS OF TROPICAL BULBS AND
ENVIRONMENTAL EFFECTS ON FLOWERING AND BULB DEVELOPMENT OF
Habranthus robustus AND Ze7,1ph/),iunhe spp.















By

CAMILA BRITO PAULA


A THESIS PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
MASTER OF SCIENCE

UNIVERSITY OF FLORIDA


2006

































Copyright 2006

by

Camila Brito Paula
































This document is dedicated to my husband and to my parents in Brazil.















ACKNOWLEDGMENTS

I thank Dr. Rick Schoellhom for giving me the opportunity of entering the "plant

world," for believing that an architect could be able to learn some science and for serving

as my major advisor for the first half of the process of my research. I thank Dr. Dennis

McConnell for playing a large role in my research from the beginning, for making my

"bulb-killing" experience much more enjoyable, for accepting the challenge of becoming

the chair of my committee in the middle of the process, and also for his enormous

patience and willingness to help me during the preparation of this thesis. I thank Dr.

Wagner Vendrame for helping me start this whole process and for much advice. I also

thank Dr. Alan Meerow for serving on my supervisory committee.

I thank Mrs. Fe Almira for her immense help with the images of this thesis, Mrs

Carolyn Bartuska for her help with experiments set ups and statistical analysis, and

Robert Weidman and his crew for all their greenhouse assistance.

I would like to thank my husband for the encouragement, the emotional and

"technical" support, for his love and friendship. And I thank my mom and dad for their

unconditional love, "long-distance" support and for giving me the educational and

emotional base that allowed me to achieve my dreams. Lastly, I would like to thank all

the friends I made in Gainesville, especially Carmen Valero-Aracama.
















TABLE OF CONTENTS

page

A C K N O W L E D G M E N T S ................................................................................................. iv

LIST OF TABLES ............. ................... ........... .. .. ............... .. vii

LIST O F FIG U R E S ......................................................... ......... .. ............. viii

A B S T R A C T .......................................... ..................................................x v

CHAPTER

1 INTRODUCTION ............... ................. ........... ................. ... .... 1

2 LITER A TU R E REV IEW ............................................................. ....................... 6

A erial O rgans and Tissues .......................................................... .............8
U underground Tissues ....................................................... ..... ........ .. 8
Taxonom y and Origin of Geophytes ........................................ ........................ 9
Geophytes Growth and Developm ent...................................... ......................... 9
D o rm a n c y ............................................................................................................. 1 0
Flow ering Process .............. .................. ....................... .............. ... 12
Anatomy and Physiology of Flower Initiation..............................................14
Factors Affecting Flower Initiation ..........................................................17
P hotoperiod .................................................................... 18
Light quality and quantity ........................................ ........................ 18
T em p eratu re ................................................. ................ 19
V ernalization .............................................. ............... ......... 20
Flower Initiation Process in Bulbous Plants..................................................21
F low er B ulb C ultiv action ..................................................................... ..................2 1
F low er B ulb F orcing ............ ...... ...................................................... .. .... ..... .. 22
Tropical Bulbs and A m aryllidaceae ........................................ ....................... 24
H ipp eastrum spp ................... ...... ......................................................... ... .... .... .. 2 5
Scadoxus m ultiflorus Blood Lily ........................................ ......... ............... 26
Agapanthus africanus African Lily ...................................... ....................... 27
Habranthus robustus and Z7,eph)i whihe, spp. Rain Lilies.................... ........ 28

3 B U L B M O R PH O L O G Y .................................................................... ..................31

Comparative Study ............................... .. ................ .............. ...3 31


v









M materials and M ethods ........................................................... .....................3 1
R results and D discussion ......................................................... ............... 33
B ulb type and size ........................................................ .... .. .......... 33
Leaf arrangem ent and m orphology ................................... .................37
F lo ra l in itia tio n ....................................................................................... 4 1
Meristematic region................................................... 48
General anatomical and morphological arrangement..............................55
Experiment 1................................... .......60
M materials and M ethods .............................................. .............................. 61
R e su lts ........................................................................................................... 6 1

4 ENVIRONMENTAL EFFECTS FERTIGATION FREQUENCY AND
FERTILIZER RATES ON FLOWERING IN Habranthus robustus AND
Z ,1 h sp p ............. .................................................................................... 6 4

E x p e rim e n t 1 ......................................................................................................... 6 7
M materials and M ethods .............................................. .............................. 67
R results and D discussion ......................................................... ............... 69
E xperim ent 2...........................................................................................84
M materials and M ethods .............................................. .............................. 84
R results and D discussion ......................................................... ............... 86

5 ENVIRONMENTAL EFFECTS LIGHT LEVELS ON FLOWERING AND
BULB DEVELOPMENT IN Habranthus robustus AND Zep,1iywiiel SPP...........93

M materials and M methods ....................................................................... ..................94
E x p e rim e n t 1 ................................................................................................. 9 4
E x p e rim e n t 2 ................................................................................................. 9 6
R esu lts an d D iscu ssion ......................................................................................... 9 7
E x p e rim e n t 1 ................................................................................................. 9 7
E x p erim ent 2 ...............................................................10 0

6 CONCLUSIONS ............................................ ..................113

LIST OF REFERENCES .......................................... .................. ..........119

BIOGRAPHICAL SKETCH ..................................... ........ ....................... 127
















LIST OF TABLES


Table page

3-1 Tropical geophytes used in the morphological studies ........................................32

3-2 Geophyte size and characteristics of plants examined..................................34

3-3 Leaf size of plants exam ined ........... ......... ...... ......... .................... ............... 39

3-4 Relationship between leaf and flower formation, and number of flowers per
inflorescence of plants exam ined .................................... .............. ................... 43

4-1 M ean bulb size and weight prior to experiment.....................................................85

5-1 Photometric readings and temperatures at three locations / treatments ...................96

5-2 Rainfall monthly summary in inches for Gainesville area in 2004 and 2005
according to the Florida Automated Weather Network .......................................97
















LIST OF FIGURES

Figure page

3-1 From left to right Zephyranthes, Habranthus, Hippeastrum, Scadoxus bulbs, and
im m ature Agapanthus rhizom e.. .......................... .............................................. 35

3-2 Sections of Zepi7, j Ii he,\ bulbs ....................................................... .............. 35

3-3 Sections of H abranthus bulbs.. ........................... ............................................... 36

3-4 Sections of H ippeastrum bulbs.. .......................... .............................................. 36

3-5 Sections of Scadoxus bulbs ............... .............. ..........................................36

3-6 Longitudinal sections of an Agapanthus rhizome.............. ...................................37

3-7 Hippeastrum leaf arrangement............................................. ..................... 39

3-8 H abranthus leaf arrange ent......... ............................................... ... ............ 39

3-9 Zepi7,ih) 1,inhei leaf arrangement...................................................... ............. 40

3-10 Agapanthus leaf arrangement...... .............................................. ............... 40

3-11 Scadoxus leaf arrange ent ................ .................................................................40

3-12 Longitudinal sections of a Hippeastrum bulb. ................................. ...............43

3-13 Sections of H abrantus bulbs .............................................................................43

3-14 Sections of Zepijh i /,wi bulbs ....................................................... .............. 44

3-15 Cross section of a Habranthus bulb .............................. ...............44

3-16 Longitudinal section of a Habranthus bulb................................... ............... 45

3-17 Habranthus bulb with three flower stalks................................................ 45

3-18 Development of a Habranthus flower............. ............... ......... ...............46

3-19 Sections of Scadoxus bulbs ............... .............. ..........................................46









3-20 Flower stalk emerged and pseudo-stem arising from a Scadoxus bulb....................47

3-21 Sections of Agapanthus rhizom es.. ............................................... .....................47

3-22 Photomicrograph of the meristematic region of a Hippeastrum bulb...................50

3-23 Photomicrograph of the meristematic region of a Hippeastrum bulb...................50

3-24 Photomicrograph of the meristematic region of a Habranthus bulb..................... 51

3-25 Photomicrograph of the meristematic region of a Habranthus bulb..................... 51

3-26 Photomicrograph of the meristematic region of a Zep,1iyittwhe, bulb....................52

3-27 Photomicrograph of the meristematic region of a Zep,1iyi ,aiwhe bulb....................52

3-28 Photomicrograph of the meristematic region of a Scadoxus bulb..........................53

3-29 Photomicrograph of the meristematic region of a Scadoxus bulb..........................53

3-30 Photomicrograph of the meristematic region of an Agapanthus rhizome..............54

3-31 Photomicrograph of the meristematic region of an Agapanthus rhizome..............54

3-32 Longitudinal section of a Hippeastrum bulb showing its general anatomy and
m orphology........................................................ ........ ........ ........... 55

3-33 Longitudinal section of a Hippeastrum bulb showing its general anatomy and
m orphology........................................................ ........ ........ ........... 55

3-34 Longitudinal section of a Zep,1il),aiiihe bulb showing its general anatomy and
m orphology ....................................................................... .. ....... ....... 56

3-35 Longitudinal section of a Zep,1 ii iite, bulb ..................................................... 56

3-36 Sections of H abranthus bulbs ............................................................................ 57

3-37 Longitudinal section of a Scadoxus bulb showing its general anatomy and
m orphology. ........... ........... ....... .. ........................... ......... 57

3-38 Longitudinal section of a Scadoxus bulb showing its general anatomy and
m orphology ........................................... ........................... 58

3-39 Longitudinal section of an Agapanthus rhizome showing its general anatomy
an d m orph ology ............................................................................... ............... 59

3-40 Longitudinal section of an Agapanthus rhizome showing its general anatomy
an d m orph ology ............................................................................... ............... 59









3-41 Number of leaves and flowers produced by Habranthus bulbs in 2005 .............. 63

3-42 Number of leaves and flowers produced by Zep,1yiinihe,\ bulbs in 2005 ..............63

4-1 Habranthus bulbs during fertigation experiment with different plastic tags in
different color distinguishing the three treatments...........................................68

4-2 Habranthus bulbs being weighed after completion of experiment.......................69

4-3 Number of flowers ofHabranthus robustus bulbs in year 2 as affected by
fertigation frequency ...................................................................... ...................74

4-4 Number of leaves of Habranthus robustus bulbs in year 1 as affected by
fertigation frequency ...................................................................... ...................74

4-5 Number of offsets ofHabranthus robustus bulbs in year 1 as affected by
fertigation frequency ...................................................................... ................... 75

4-6 Bulb size of Habranthus robustus in year 1 as affected by fertigation frequency...75

4-7 Total weight ofHabranthus robustus bulbs in year 1 as affected by fertigation
fre q u e n cy ......................................................................... 7 5

4-8 Bulb weight of Habranthus robustus in year 1 as affected by fertigation
fre q u e n cy ......................................................................... 7 6

4-9 Number of flower buds of Habranthus robustus bulbs in year 1 as affected by
fertigation frequency ...................................................................... ...................76

4-10 Number of leaves of Habranthus robustus bulbs in year 2 as affected by
fertigation frequency ...................................................................... ...................76

4-11 Number of offsets ofHabranthus robustus bulbs in year 2 as affected by
fertigation frequency ...................................................................... ...................77

4-12 Bulb size of Habranthus robustus in year 2 as affected by fertigation frequency...77

4-13 Total bulb weight of Habranthus robustus in year 2 as affected by fertigation
fre q u e n cy ......................................................................... 7 7

4-14 Bulb weight of Habranthus robustus in year 2 as affected by fertigation
fre q u e n cy ......................................................................... 7 8

4-15 Number of flower buds of Habranthus robustus bulbs in year 2 as affected by
fertigation frequency ...................................................................... ................... 78

4-16 Total number of flowers on Habranthus and Zep,1 iyiiie1 \ bulbs from July to
D ecem ber 2004 .......................................................................79









4-17 Number of flowers of Ze7,y'l)-1/,ihe spp. bulbs in year 1 as affected by
fertig action frequ en cy ........................................................................ .................. 79

4-18 Number of flowers of Ze7,y'l)-1/1ihe spp. bulbs in year 2 as affected by
fertigation frequency ...................................................................... ................... 80

4-19 Number of leaves of Ze7,1'/ji/,he, spp. bulbs in year 1 as affected by fertigation
fre q u e n cy ......................................................................... 8 0

4-20 Number of offsets of Ze7,y'1,l)-i/wh spp. bulbs in year 1 as affected by fertigation
frequency. .............................................................. ..... .......... 81

4-21 Bulb size of Ze7,y h)/1/i,,/ spp. in year 1 as affected by fertigation frequency. ......81

4-22 Total fresh weight of Ze7,y'1)l-i/ih spp. bulbs in year 1 as affected by fertigation
frequency. ................ ........ .......................................................... 81

4-23 Bulb weight of Ze7,y')1/iwhe, spp. in year 1 as affected by fertigation frequency...82

4-24 Number of flower buds of Ze7,1,h)Irl /wi spp. bulbs in year 1 as affected by
fertigation frequency ...................................................................... ................... 82

4-25 Number of leaves of Z7,1iy'l-i/,he, spp. bulbs in year 2 as affected by fertigation
freq u en cy .............................................................................................. 8 2

4-26 Number of offsets of Ze',1,yl)-i/he, spp. bulbs in year 2 as affected by fertigation
fre q u e n cy ......................................................................... 8 3

4-27 Bulb size of Zep7,10l)y ,,he spp. in year 2 as affected by fertigation frequency. ......83

4-28 Total fresh weight of Ze7,iy')l-i/he, spp. bulbs in year 2 as affected by fertigation
fre q u e n cy ......................................................................... 8 3

4-29 Bulb weight of Ze7,y'l)-1i1.ih spp. in year 2 as affected by fertigation frequency...84

4-30 Number of flower buds of Ze7,1,h)Irl /wi spp. bulbs in year 2 as affected by
fertig action frequ en cy ........................................................................ .................. 84

4-31 Data points, regression lines, equations and coefficient of determination of
number of leaves of Habranthus ................. .. ......................... ............... 88

4-32 Data points, regression lines, equations and coefficient of determination of
number of offsets of Habranthus. ..........................................................................88

4-33 Data points, regression lines, equations and coefficient of determination of bulb
size of Habranthus. ............................ ...... ..... ........... ...... 89

4-34 Data points, regression lines, equations and coefficient of determination of total
fresh bulb w eight ofH abranthus .......................................................................... 89









4-35 Data points, regression lines, equations and coefficient of determination of fresh
bulb weight of Habranthus .................................. ......................................89

4-36 Data points, regression lines, equations and coefficient of determination of
number of flower buds of Habranthus............................................. ............... 90

4-37 Data points, regression lines, equations and coefficient of determination of
number of leaves of Z7 1 h 1....................... ............ ................. .......... 90

4-38 Data points, regression lines, equations and coefficient of determination of
number of offsets of Z7,1, h, ....................................... ................... ..90

4-39 Data points, regression lines, equations and coefficient of determination of bulb
size of Zephyranthes .................. ............................ .. ...... ... ........ .... 91

4-40 Data points, regression lines, equations and coefficient of determination of total
fresh bulb w eight of Zep7,1 i he, ......................................................................... 91

4-41 Data points, regression lines, equations and coefficient of determination of fresh
bulb w eight of Zep7,1/) iiw he, .................. ............................................... .. ... 91

4-42 Data points, regression lines, equations and coefficient of determination of fresh
bulb w eight of Z 7 j, ) i he,' ................................................................................ 92

5-1 Plants under three different treatments full sun ....................................... .......... 96

5-2 Number of flowers ofHabranthus robustus bulbs in year 2 as affected by light
le v e ls .......................................................................... ................ 1 0 2

5-3 Number of leaves ofHabranthus robustus bulbs in year 1 as affected by light
le v e ls .......................................................................... ................ 1 0 3

5-4 Number of offsets ofHabranthus robustus bulbs in year 1 as affected by light
le v e ls ...................................................................................... 1 0 3

5-5 Bulb size of Habranthus robustus in year 1 as affected by light levels................ 103

5-6 Total weight of leaves ofHabranthus robustus bulbs in year 1 as affected by
light levels. .......................................... ........................... 104

5-7 Bulb weight ofHabranthus robustus in year 1 as affected by light levels ..........104

5-8 Number of flower buds of Habranthus robustus bulbs in year 1 as affected by
light levels. .......................................... ........................... 104

5-9 Root condition of Habranthus bulbs, under different treatments, after
com pletion of experim ent 1 ............... ........................................................... 105









5-10 Number of leaves ofHabranthus robustus bulbs in year 2 as affected by light
le v e ls ................................................... .................... ................ 1 0 5

5-11 Number of offsets ofHabranthus robustus bulbs in year 2 as affected by light
lev e ls ...................................... .................................................. 1 0 5

5-12 Bulb size of Habranthus robustus in year 2 as affected by light levels...............06

5-13 Total weight ofHabranthus robustus bulbs in year 2 as affected by light levels.. 106

5-14 Bulb weight ofHabranthus robustus in year 2 as affected by light levels ..........106

5-15 Number of flower buds of Habranthus robustus bulbs in year 2 as affected by
light levels. .......................................... ........................... 107

5-16 Number of flowers of Ze7,y'1)li/iwh spp. bulbs in year 1 as affected by light
levels ............. ..... .... ......... ............ ............... 107

5-17 Number of flowers of Ze7,y')1l-i/ih spp. bulbs in year 2 as affected by light
levels ............. ..... .... ......... ............ ............... 108

5-18 Number of leaves of Ze7,y'p/1)liih spp. bulbs in year 1 as affected by light
lev els ............... ......... ................................ .............................. 10 8

5-19 Number of offsets of Zepy',1/lihe, spp. bulbs in year 1 as affected by light
levels ............... ...... ............................................... ........... 108

5-20 Bulb size of Zep7y,1/) i e, spp. in year 1 as affected by light levels. ..................109

5-21 Total weight of Ze7,py1imwhi/e spp. bulbs in year 1 as affected by light levels. .....109

5-22 Bulb weight of Zep7',1w1iihe, spp. in year 1 as affected by light levels................. 109

5-23 Number of flower buds of Zepy7,1t),i/wh spp. bulbs in year 1 as affected by light
lev e ls ......... .. ........................................................................ .. ........ ... 1 1 0

5-24 Number of leaves of Ze7,1pl-ii/he, spp. bulbs in year 2 as affected by light
levels ........... .. ...... ............ ......... ........... 110

5-25 Number of offsets of Ze',1p /l)-r he spp. bulbs in year 2 as affected by light
levels ................ ...... ............................................... ........... 110

5-26 Bulb size of Zep7y,1/) i/w spp. in year 2 as affected by light levels. ..................111

5-27 Total weight of Ze7,iy 1,) im, spp. bulbs in year 2 as affected by light levels. .....111

5-28 Bulb weight of Zep7,1y'i-in/w spp. in year 2 as affected by light levels ................11









5-29 Number of flower buds of Ze7ph,1/It/.he spp. bulbs in year 2 as affected by light
lev e ls ...................................... .................................................. 1 12















Abstract of Thesis Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Master of Science

MORPHOLOGICAL ANALYSIS OF TROPICAL BULBS AND
ENVIRONMENTAL EFFECTS ON FLOWERING AND BULB DEVELOPMENT OF
Habranthus robustus AND Ze7,1ylu-iiihe spp.

By

Camila Brito Paula

May 2006

Chair: Dennis B. McConnell
Major Department: Horticultural Sciences

Morphologies of Hippeastrum x hybridum, Habranthus robustus, Ze7,1'1i i uin1h

spp, Scadoxus multiflorus, and Agapanthus africanus were compared and contrasted. The

first three species were true tunicate bulbs with similar shapes and distinct sizes, leaves

emerged alternately from the center of the bulb, flower buds were formed alternately at

the apical meristem (in a line in bulb cross section), and there were four leaves between

each flower formed. Scadoxus was a true tunicate bulb with a thick rhizomatous structure

at the base, thick scales, leaves arising in a pseudostem and flower buds formed centrally

at the apical meristem. Its size is equivalent to Hippeastrum bulbs. Agapanthus is a

rhizome with leaves arising alternately from lateral meristems, and flower buds formed

centrally at the apical meristem.

Habranthus and Zp7,1hl1yiiu1he had distinct patterns of leaf production. Habranthus

bulbs had fewer leaves emerging during flowering compared to 7eph,1yi/,inhe,, which had

a greater number of leaves throughout the entire year.









The responses of Habranthus robustus and Ze7,10/)yiiwh spp. to fertigation

frequencies and fertilizer rates were examined. Experiment 1 tested fertigation

frequencies of twice a week, once a week and every other week in both species.

Regimens of once or twice a week were effective for flowering of both species. All bulb

development factors investigated increased as fertigation increased in Habranthus and

Zephyranthes, except for number of offsets on both species and number of leaves in

ZLT'/'1/I tlI//l"'\

Experiment 2 tested bulb development responses when both species were fertilized

with 20-10-20 at rates of 0, 75, 150, 300 ppm N. Habranthus bulbs treated with 75 and

150 ppm N had the greatest number of leaves and flower buds, and were larger and

heavier, but bulbs treated with 300 ppm had more offsets. Ze7,1yiiniwhe bulbs treated

with 150 ppm N had the greatest number of leaves, offsets and flower buds; bulbs treated

with 300 ppm N were largest. Bulb weight was similar in bulbs treated with 75, 150 or

300 ppm N.

The responses of Habranthus robustus and Ze7,1',)-i/iwh spp. to light levels of full

sun, 30% and 60% shade were examined. Results demonstrated that both Habranthus and

Ze7,1'l)I -ihe, flowered for a longer period under full sun and 30% shade than 60% shade

but plants had more flowers under 30% shade. All bulb development factors investigated

increased as light level increased in both species.

From this study it was concluded that preferred conditions for both species were

fertigation of twice a week using fertilizer rate of 150 ppm N under full sun.














CHAPTER 1
INTRODUCTION

Bulbous plants are plants with a self-contained and highly developed food-storage

mechanism that allows them to live underground. These plants are found all around the

world; however the principal areas that are natural common habitats of a high percentage

of bulbous plants are between 23 North and 45South latitudes (Du Plessis and Duncan,

1989) in the Mediterranean, South Africa, Middle East and the Pacific seaboard of North

and South America (Bryan, 1989).

An enormous number of flower bulb genera and species are found in nature, and

they provide material for a wide range of potential ornamental use. The diversity of

flower color, form, size, habitat and desirable growing conditions of bulbous plants rivals

most other forms of vegetation. They can be used in the landscape, as borders and in

flower beds, in containers outdoor and indoor and as cut flowers in indoor arrangements.

Historically bulbs were introduced in Europe almost 400 years ago (Bryan, 1989).

Agapanthus africanus, Dur et. Schinz (1893) from South Africa was introduced in

England in 1629. Amaryllis bealladona was introduced to Europe in the late 17th century

(Traub, 1958a) and the genus Hippeastrum in the beginning of the following century

(Tjaden, 1979).

Although no registries exist, it is likely that Amaryllis belladonna was originally

collected in South Africa at the time of the spice and slave trade late 15th century and

early 16th century, possibly due to its abundance, coastal distribution, floral attributes

and commercial value. The slave trade and sugar cane production played a major role in









the distribution ofA. belladona around Europe explaining why it is still commonly found

in the Canary Islands, Madeira, Spain, Italy, and the Azores, which were main sugar cane

growing areas.

During the 1500's, European plant explorers searched the world recording new

species of plants using botanical illustrations. One of the earliest and best known

illustrations in Europe was of Tulipa bononiensis, a species of bulb that would become

extensively used, appreciated and commercialized. The Dutch botanist Carolus Clusius,

head of the University of Leiden's Hortus Botanicus, the first botanical garden in

Western Europe, received several tulip bulbs and seeds from Turkey and started a

collection at the end of the 16th century. Clusius bred the tulips and produced new color

variations, but he was mostly interested in its scientific importance and possibly

medicinal uses for the bulbs. However, people in Holland were already interested in the

flowers as money-makers for the developing ornamental floral trade, and some of

Clusius' tulips were stolen from his gardens. That was the beginning of the famous

"Tulipomania" (Cremers, 1973).

Throughout the early 1600's tulips were widely traded in the market and their

prices were extremely high. In 1624, one tulip type was sold for 3000 guilders per bulb,

the equivalent of US $1,500.00 nowadays. Since then the Dutch have built one of the best

organized bulb production and export businesses in the world. In 2001, over nine billion

flower bulbs were produced in Holland, and about 80% were exported. According to the

Netherlands Flower Bulb Information Center, the United States is the biggest importer of

Dutch bulbs.









Tulips and other spring blooming bulbs are still closely associated with the

Netherlands; however bulb production is not exclusive to that area. The world flower

bulb industry produces a wide variety of high quality bulbs adapted to many climatic

zones that are marketed either as cut flowers, flowering potted plants or landscape plants.

Today's bulb industry is firmly established in areas of the world where the growers can

duplicate by various cultural practices the environmental factors required for floral

initiation and development. The demand for bulb crops is high, and the potential exists

for greater production and sales of these crops (Johnson et al., 1995).

Florida is the second largest floriculture producing state in the US with over $650

million in sales according to USDA, making floriculture a vital part of Florida's

agricultural economy. Its hot and humid summers and rare frost occurrences during

winter provide an ideal scenario for a varied selection of bulbous plants.

The tropical bulb market is practically non-existent in Florida due to production

and market problems. Many tropical bulbs grow slowly, some have viral problems, and

many consumers are not sure how to grow them. However, there is a large selection of

bulbs suitable for Florida and other regions in the US in USDA Hardiness Zones 9, 10

and 11 that produce great spring and summer color. South Florida's climate is favorable

for growing most of the tropical and subtropical bulbs. Examples of suitable tropical

bulbs for most of Florida include: African Lily, or Lily of the Nile (Agapanthus

africanus), Amaryllis (Hippeastrum spp.), Spring Calla Lily (Zantedeschia spp.),

Peruvian Daffodil (Ismene narcissiflora), and Rain Lilies (Habranthus and Zep7 in,1,/n/hw,)

(Black et al., 1990).









It is important to increase our research concerning flower bulbs, particularly

subtropical and tropical bulbs, in order to identify and understand cultural factors of as

many genera as possible. This will increase the number of bulbous plants that can be

economically produced for the ornamental market. However, to be economically

competitive, these plants should be profitable to produce and market, multiply under a

wide variety of soil and climatic conditions, adapt to mechanical handling, tolerate air

pollution, resist periods of drought, and tolerate low to moderate nutritional status.

The present study addressed some of these issues and was designed to provide

growers with improved methodology and guidelines for the commercial production of

two genera of tropical bulbs Habranthus robustus (Herb Lodd. 1831) and

Ze7,1)1/iuiwhe, spp. (Herb., 1821). The genera Habranthus and Ze',1-iawiie, belong to the

Amaryllidaceae family and are known as Rain Lilies since they bloom several times

during a season, usually following rainfalls. Limited information is available in the

horticultural literature on cultural practices and commercial production of these two

genera (especially of Habranthus) and their ability to adapt and develop under Florida's

climatic conditions. Increased knowledge of these factors could result in the plants

becoming important commercial crops in Florida.

This study was designed to determine the effect of environmental factors of light

levels, drought stress, and the cultural practices of irrigation frequency and fertilization

on flowering response of Habranthus robustus and Ze7,10yl, i1whe spp. Specific objectives

of this study were 1) to perform morphological evaluations on Habranthus robustus and

Ze7',11i waihe, spp. bulbs, comparing those with other species of tropical bulbous plants;

2) to evaluate environmental effects on flowering responses for Habranthus robustus and






5


Ze7,yl ranihe\ spp.; and 3) to develop commercial production and cultural information for

Habranthus robustus and Ze7h,1)I/, // iah spp.














CHAPTER 2
LITERATURE REVIEW

Geophyte, from the Greek geo for earth and phyton for plant; is a term that refers to

plant species with specialized underground storage organs that accumulate food reserves,

nutrients and moisture for seasonal growth and development. This group of plants

includes both monocotyledonous and dicotyledonous species (Bryan, 1989) and is

collectively referred to as "flower bulbs" (Halevy, 1990; Rees, 1985).

Geophytes can be separated into four groups true bulbs, corms, tubers and

rhizomes. Although they are morphologically different, the underground portions of all

types of geophytes perform the same basic function storage. A key factor used to

classify geophytes is the precise origin and nature of the tissue that serves as the primary

storage tissue (De Hertogh and Le Nard, 1993). Despite morphological differences, all

geophytes share a common characteristic: they have a self-contained, highly developed

food-storage mechanism.

True bulbs have a shortened stem, with a basal plate, one or more apical meristems,

enclosed flower buds, adventitious roots initials, several layers of fleshy scales and a

protective tunic that envelops the bulb. The scales are modified leaves (enlarged leaf

bases) and function as the primary storage tissue in true bulbs. The tunic protects the bulb

from drying and mechanical injuries (De Hertogh and Le Nard, 1993).

Bulbs can be either tunicate or non-tunicate depending on the origin of their scales.

Concentric layers of scales form tunicate bulbs, such as in Tulipa, Hippeastrum, and









Narcissus. Non-tunicate bulbs do not develop concentric or scaly layers of scales, such as

Lilium (Black et al., 1990).

Small underground bulbs are either called bulblets or offsets, if they occur at the

periphery of the mother bulb. Small aerial bulbs that occur in either the leaf axils or in the

floral parts are called bulbils. Most true bulbs are monocotyledonous, such the genera

Allium, Amaryllis, Haemanthus, Habranthus, Lilium, Nerine, Tulipa and Z7,1yii ,un1e\

(Bryan, 1989).

Rhizomes are horizontal, thickened, branching storage stems which grow below or

along the surface of the soil. Typically, shoots (above ground) and roots (on the lower

surface) arise at right angles from the cross stem. They are monocotyledonous

(Agapanthus, Canna and Clivia), and dicotyledonous (some Anemone and Achimenes).

Corms are solid masses of stem tissue with an enlarged basal plate, distinct nodes

and intemodes, and adventitious root initials, enclosed by several dry, papery scale-like

leaves. They are protected against injury and water loss by dry leaf bases that are similar

to the tunic that encloses true bulbs. Corms are easily distinguished from true bulbs by

the absence of visible storage rings when crossly sectioned and by having the basal plate

as the primary storage organ. Small corms are called cormlets or cormels. Most corms

reproduce by annual replacement and are monocotyledonous; examples include Crocus,

Freesia and Gladiolus (Bryan, 1989).

Tubers consist basically of enlarged underground stem tissue with a root primodia

developing basally and one or more apical shoot meristems (buds) on the underground

stem. These buds (also called eyes) arise from the nodes and they are arranged in the

same spiral pattern characteristic as buds on an aerial stem. Tubers are covered with a









tough skin and do not have scales as in true bulbs. Another distinction from true bulbs is

the absence of a basal plate and protective tunic. They occur in both monocotyledonous

(Caladium, Gloriosa and some Zantedeschia) and dicotyledonous (Anemone and

Eranthis) genera (Bryan, 1989).

Aerial Organs and Tissues

Geophytes exhibit a wide variety of flowers, stems and leaves (Bryan, 1989). Their

flowers can be single, double, semi-double or multiflowered, depending on the number of

petals. According to Du Plessis and Duncan (1989) they can be hysteranthous (leaves

appear after flowers), proteranthous (leaves appear before flowers) or synanthous (leaves

and flowers appear simultaneously). Floral development can be determinate (number of

flowers do not increase after first flower is opened) or indeterminate (number of flowers

can increase after first flower is opened). Stems can be leafed or leafless. Plants can be

multi-stemmed or single-stemmed. Additionally, geophytes can be classified as either

evergreen or deciduous according to their leaf persistence throughout the year.

Underground Tissues

Three morphological characteristics are commonly used to describe flowering bulb

root systems: branching habit, presence or absence of root hairs and contractile root habit.

Although most higher plants produce branched root systems (Whittington, 1969), several

ornamental flowering bulbs do not. No branching has been observed for several genera

such as Allium, Crocus, Hyacinthus, Muscari, Narcissus, and Tulipa (Kawa and De

Hertogh, 1992). It is generally assumed that all plant roots have root hairs, but this is not

true for some ornamental flower bulbs such as Crocusflavus (De Munk and De Rooy,

1971).









Contractile roots play a major role in positioning bulb crowns and storage organs at

a proper soil depth for optimal growth and survival of the species. Corms and true bulbs

commonly reposition themselves in the ground via contractile roots. This mobility of the

underground plant organs occurs because of the activity of the contractile roots in

response to light and temperature (De Hertogh and Le Nard, 1993). Of the bulbs used in

their study, Theron and De Hertogh (2001) reported that Hippeastrum, Agapanthus and

Z.7'p,1/1i lhe have contractile roots, while Scadoxus did not have contractile roots.

Habranthus was not used in their study.

Rhizomes secure an optimal position by the growing activity of their shoots tips, as

reported by Ptitz (1998) who found that Hemerocallisfulva rhizomes were able to adjust

their own soil depth by elongating the root axis and not via root contraction.

Taxonomy and Origin of Geophytes

The three most important plant families containing geophytes are Amaryllidaceae,

Iridaceae and Liliaceae. Most tropical bulbs, and those used in this study, belong to the

family Amaryllidaceae. Bryan (1989) summarized the origin of many geophytes and

stated that they mainly occur between the 23 to 450 North and South latitudes, this

includes the Mediterranean, South Africa, Middle East and the Pacific seaboard of North

and South America.

Geophytes Growth and Development

Geophytes must reach a certain physiological stage before they are capable of

flowering; this can take less than a year or as long as six years (De Hertogh and Le Nard,

1993). In several species the ability to flower is directly related to the size of the

geophytic organ, which varies from species to species. Hippeastrum for example, flowers

when bulbs reach a circumference of approximately 20 cm (6 cm in diameter), Scadoxus









flowers when the geophytic organ circumference is 15 cm (4.5 cm in diameter), while

Eucharis bulbs need to attain a circumference of 3 to 5 cm (1 cm in diameter) before

flowering occurs (Theron and De Hertogh, 2001).

A number of factors affect growth, development, and flowering in bulbs. These

include their native habitat and related microclimate parameters such as temperature

range, rainfall, sun irradiation, photoperiod, altitude, moisture, soil type and nutritional

status. With the exception of some equatorial areas that have fairly uniform

environments, geophytes are often exposed to a wide range of climatic conditions during

their growth cycle (De Hertogh and Le Nard, 1993).

Among environmental factors that affect bulb growth and development, the most

important ones are temperature, light and moisture. These three factors are manipulated

to force bulbs, since they act directly on rooting, flower development, shoot elongation,

and bulbing (bulb elongation). Temperature is the major external factor that controls

growth, development, flowering, dormancy and physiological maturity of bulbs. Light

intensity and photoperiod also affect several physiological processes, such as

photosynthesis, flower abortion and abscission (Bryan, 1989).

Dormancy

Bulbs have developed mechanisms to survive seasonal changes in climatic

conditions such as low or high temperatures and drought. Under adverse conditions many

bulbs enter a dormant period, in which they do not exhibit any visible external growth. Le

Nard (1983) stated that the period of dormancy mainly corresponded to the period of

bulbing, when the bulb enlarged. However, De Hertogh and Le Nard (1993) defined it as

a complex and dynamic physiological, morphological and biochemical state during which









there are no apparent external changes or growth. Internally, however, many

physiological and/or morphological events may occur.

Roughly, species that go dormant are deciduous as they loose their leaves during

adverse periods (winter or seasonal dry periods) and those that do not go dormant are

evergreen (they keep their leaves all year round). Examples of evergreen bulb genera

include Agapanthus, Amaryllis and Clivia. Different genera respond differently to this

period of dormancy and have also distinct requirements to break dormancy. Yet,

temperature and moisture are the two major factors used to affect bulb dormancy (Le

Nard, 1983).

Some temperate zone bulb species, such as Tulipa, Daffodil and Hyacinth, have

cold dormancy requirements; which means they need a critical number of chilling hours

in order to bloom the following season (Kamenetsky, 2004). That does not occur in

Florida even in the northern areas of the state as most winters are not sufficiently cold.

Consequently, temperate zone bulbs are not used in the landscapes, except as annuals, in

these areas. Other species, such as Caladium and Zantedeschia, have warm dormancy

requirements, which means they need a period of dry and warm temperature in order to

bloom next season (De Hertogh and Le Nard, 1993). North and Central Florida's winters

are usually not sufficiently dry or warm for some of these species; however, South

Florida's winters are suitable for some of the species.

The existence of a rest or dormant period is very convenient for horticultural

practices because it facilitates the handling, storage and transportation of these bulbs. The

species used in this study, Habranthus spp. and 7Zephyi ,hi.en spp, go dormant during the

cold season and require very little water during this period. However, not all bulb crops









go through a dormant period. Hippeastrum bulbs, for example, do not require a rest

period in order to flower, since bulbs can produce inflorescences annually in a

greenhouse under conditions of continued irrigation (Haynes et. al, 2001).

Flowering Process

Plants continue to generate new organs even after their embryonic phase, unlike

animals. Undifferentiated cells called meristematic cells are responsible for the formation

of new organs. The organ produced by the shoot meristem during its post-embryonic

phase will depend on the phase of the plant's life cycle, which can be characterized as:

juvenile vegetative phase, adult vegetative phase and reproductive phase. During the

juvenile phase, the shoot meristem initiates stems, true leaves and axilary buds; and

during the adult phase, the shoot meristem can form inflorescences which contain sexual

organs. In some plants reproduction is the last of the shoot's phases; in other plants

vegetative growth begins on lateral meristems when the apex becomes reproductive. In

some other plants the apical meristem remains vegetative and lateral meristems generate

reproductive structures (Poethig, 1990).

The transition from producing one organ to another, which is known as phase

change, can be either gradual or abrupt. This process, called flower initiation, marks the

end of vegetative growth and is a major determinant of plant reproductive success

(Poethig 1990).

The flowering process involves five successive stages: induction, initiation,

organogenesis, maturation (growth of the floral parts), and anthesis (De Hertogh and Le

Nard, 1993). Important aspects of this process are: 1) the signals received by the plant

that instigate the process, 2) their transport to the shoot apex, and 3) the changes in the

shoot apex during floral differentiation (Evans, 1993).









The successive steps of the flowering process are controlled by several factors and

each takes place in a determined period of the growth cycle. Controlling these processes

in bulbous plants, as well as other horticultural crops, can promote or retard flowering,

prevent flowering or induce flower abortion (De Hertogh and Le Nard, 1993). Promoting

or retarding flowering permits out of season and/or year-round commercial flower

production. Preventing flowering is necessary for the production of some bulbs, such as

Iris hollandica. Inducing flower abortion can promote bulb development in some bulb

species such as Tulipa and Hippeastrum. The flowering process can be controlled by

applying specific treatments (temperature, moisture, light, or plant growth regulators) to

the bulb. However, precise knowledge of bulb periodicity is essential to control flowering

(Bryan, 1989).

Flower initiation takes place at different times of the year and at different stages in

the development of bulbs. Seven different types of flower initiation have been identified

in commercially grown bulbous plants (Hartsema, 1961):

1. Flowers are formed during spring or early summer of the year preceding the one in
which they flower (Narcissus, Galanthus, Leucojun and Convallaria).

2. Flowers are formed after the end of the assimilation period (Crocus, Hyacinthus,
Iris reticulata and Tulipa).

3. Flowers are formed some time after new growth matures, in winter or early spring
(most Iris spp).

4. Flowers are formed during the storage period and complete development after
planting (Begonia tuberosa, Dahlia and Lilium).

5. Flowers are formed after replanting in spring (Galdiolus, Anemone and Freesia).

6. Flowers are formed more than a year before flowering (Amaryllis belladonna and
Nerine sarniensis).









7. Flower formation occurs alternatively with leaf formation during the whole
assimilation period (Z7eph)i,1iwnh,\ and Habranthus). In this case, both young
developing flower buds and one year-old flower buds are present in the bulb.

Anatomy and Physiology of Flower Initiation

All the aerial parts of a plant (excluding the cotyledons) are produced by the shoot

apical meristem. This meristem is formed during embryogenesis and comprises a group

of undifferentiated cells that generate different organs and structures such as stems,

leaves and flower. The root system of a plant is produced by the root meristem (Poethig

1990).

The shoot apex meristem at a certain point in the growth cycle undergoes a phase

change, caused by floral induction in response to exogenous (such as daylength, nutrient

status, or temperature) and endogenous (related to internal factors such as plant age and

metabolic status) factors, and becomes reproductive producing flower buds instead of

leaves. This transition is not irreversible in plants and in some species under certain

environmental conditions, leafy shoot formation occurs after flower formation in a

phenomenon known as inflorescence reversion (Pouteau et al. 1997).

Three different models have attempted to describe the control of flowering in

plants. The first model is known as the "florigen concept" (Evans, 1971) which suggested

that substances or signals are transferred across grafts between reproductive "donor"

shoots and vegetative "recipients", and that a flower-promoting hormone called florigen

transports them to the shoot apex via the phloem. Chailahjan (1937) suggested that the

florigen hormone was produced in leaves under favorable photoperiodic conditions. This

model could not be proved, which lead to a second model called nutrient diversion

hypothesis. This suggested that external conditions could raise the amount of assimilates

moved to the apical meristem, which were responsible for flower induction (Bernier,









1988). A third model, called "multifactorial control model", proposed that several

promoters and inhibitors (including phytohormones and assimilates) were involved in

flowering induction; and that flowering would only occur if these were present in the

apical meristem in proper concentrations and specific time intervals (Evans, 1995).

Genetic investigations of flowering in several species support the assumption that

several factors act together to control flowering. Most of these studies have been done in

Arabidopsis, and have provided recent advances in better understanding of this process

(Guo et al, 1998; Koornneef et al., 1998; Simpson, 2002).

A number of genes are involved in flower initiation, such as the flowering-time,

which integrates the signals and act as promoters or repressors of flowering, the

meristem-identity, which determines the fate of newly formed primordia (shoot/leaf-, or a

flower-primordium), and the organ-identity, which directs the formation of various

flower parts (reviewed in Yanofsky, 1995). Peeters and Koornneef (1996) have identified

several genes that control the timing of the transition from leaf production to flower

production by mutant analysis. There is still much to be learned about the functions of

these different genes, however, it has been suggested that some of these flowering-time

genes are involved in the partitioning or metabolism/sensing of compounds that might

play a part in the endogenous plant signalss, such as the plant hormone gibberellin, and

sucrose concentrations (Nilsson et al. 1998).

There is a small group of plants that can be induced to flower artificially by the

application of chemicals. Manochai (2005) tested the application of potassium chlorate

(KC103) to either the root or the foliage ofDimocarpus longan and obtained successful

results at all times of the year.









During flower induction and early flower development the apical meristem

undergoes several morphological alterations (Tooke and Battey, 2003). Changes in local

forces at the surface of the shoot apex have been proposed to play an important role in

organogenesis (Fleming et al., 1997), which is the step that follows initiation in the

flowering process. Albrechtova et al. (2004) examined the role of these local forces

during photoperiodic flower induction in Chenopodium rubrum, by measuring the

changes in shape of the apical dome, and changes in cell wall properties. The results

obtained confirmed that early changes at the surface of the apical meristem affect the

process of floral transition. The relative size of the apical meristem has also been

proposed as an important factor in the developmental (vegetative/reproductive) switch.

After a signal is received by the plant and transferred to the shoot apex, the

meristematic cells begin the process of organogenesis, the formation of tissues by cell

differentiation. These tissues mature forming the floral organs. Different species have

different mechanisms with which they control the development of their flower organs

(stamens, carpels, etc). Most species develop their carpels and stamens simultaneously,

but in some species the male flower organs grow first (these are referred to as

protandrous flowers) and in others the female organs grow first (and these are referred to

as protogynous flowers). A few species like pumpkin (Cucurbitapepo) develop male

flowers first, next hermaphroditic ones, finally female and at last parthenocarpous

flowers (lacking an embryo) that produce seeds agamically, not involving the fusion of

male and female gametes, as occurs in sexual reproduction (Evans, 1993).

A flower is considered a modified stem with shortened intemodes and bearing

modified leaves at the nodes. In essence, a flower is structured as a modified shoot or axis









with an apical meristem that does not continue to grow. A typical flower contains a

pedicel (stem) with a torus or receptacle at the end. The four main parts (or whorls) of a

flower sit on the receptacle, and they are: the calyx (group of sepals); corolla (group of

petals); the androecium (male organs filaments and anthers); and the gynoecium

(female organs stigma, style, ovary and ovule). Plant species show a wide variety of

modifications from this plan (Eames, 1961).

The basic function of flowers is to mediate the union of male and female gametes,

in the process called pollination. The period of time during which the flower is fully

expanded, functional, and thus allow the pollination process can take place is called

anthesis, and it can vary from few hours to weeks. During the maturation phase, most

assimilates obtained by the plant are used to develop the floral parts. As soon as the

flower is fully expanded and opened, assimilates start to be mobilized and used for seed

development, and that is when the process of flower senescence begins (Poethig, 1990).

Factors Affecting Flower Initiation

In some species, the timing of flowering is primarily influenced by environmental

factors, which determine the time of year and/or growth conditions that are favorable for

sexual reproduction and seed maturation. These factors include photoperiod (day length),

light quality (wave length), light quantity (photon flux density), temperature,

vernalization (exposure to a defined number of hours below a critical temperature), and

nutrient and water availability (Halevy, 1990).

Some species are not very sensitive to environmental variables and these appear to

flower in response to endogenous factors such as plant size (bulbs are an example of that)

or number of vegetative nodes. Flowering can also be induced by stresses such as nutrient

deficiency, drought, and overcrowding. This response enables the plant to produce seeds,









which lay dormant until the return of favorable environmental factors (Levy and Dean,

1990).

Photoperiod

Photoperiod is defined as the length of time a plant is exposed to light and/or

darkness within a 24-hour period. Flower induction, initiation and/or development of

many plant species are greatly affected by photoperiod and can be synchronized during

the year by manipulating night length (Garner and Allard, 1920). According to Thomas

and Vince-Prue (1997), plants with photoperiodic flowering responses are divided into

five groups according to the amount of light/darkness they require: short-day (SD) plants,

long-day (LD) plants, day-neutral (DN) plants, intermediate-day plants, and

ambiphotoperiodic-day plants. Short- and long-day plants can also be subdivided into:

obligate or qualitative, and facultative or quantitative. Photoperiodic responses are often

interspecific and can vary within cultivars of a species (Martson and Erwin, 2005). In

most plants the amount of darkness is what determines initiation; a night break (or

addition of extra light during the dark period) can inhibit or promote flowering (Evans

1993).

Rees (1985) reported that Hippeastrum bulbs have no response to photoperiod

regarding flowering. However, De Hertogh and Gallitano (2000) determined that

photoperiod, as well as temperature, affect leaf size in the 'Apple Blossom' cultivar of

Hippeastrum.

Light quality and quantity

There are two different plant behaviors related to light quality: light dominance and

dark dominance. Light dominants are plants that are susceptible to changes in the spectral









distribution during the light period. Dark dominants are plants that need uninterrupted

dark periods in order to flower (Fankhauser and Chory, 1997).

Flower initiation and time to flower in LD plants are related to light quality and to

the timing of light treatments in the photoperiod (Thomas and Vince-Prue, 1997). Pringer

and Cathey (1960) examined the effects of different types of light on Petunia flowering

and concluded that they flower 2 to 3 weeks earlier if exposed to incandescent light

(containing red and far-red light) than when exposed to fluorescent light (which has no

far-red light).

Red light appears to be effective earlier in the photoperiod for LD plants, while far-

red light seems to be important in the flower inductive response later in the photoperiod

for LD plants, which according to Thomas and Vince-Prue (1997) is different from SD

plants. Studies made by Vince (1965) in Lolium temulentum showed that red light used

for 8 hr at the beginning or at the end of the photoperiod induced flowering, but when it

was used at the beginning it promotes stronger flowering.

Temperature

In bulbous plants, temperature is the major external factor controlling growth,

development and flowering (promoting or delaying it). Temperature also affects bulb

dormancy in some genera and the physiological maturity of the bulbs (Bryan, 1989).

Doorduin and Verkerke (2002) investigated bud development and flowering of

Hippeastrum under 15 to 250C and observed that at higher temperatures larger bulbs with

more leaves developed, but the percentage of bulb dry matter decreased. In Scadoxus,

temperature affects not only flowering but also bulb development. The optimal

temperature to overcome dormancy of Scadoxus bulbs is 10 to 150C, which is low

enough to injure several other species (Bryan, 1989).









Flower formation does not occur in Amaryllis belladonna at 90, 130 or 31C, and

the optimal temperature for flower initiation on this species appears to be 17C. However,

for non-planted Amaryllis belladonna bulbs, the optimal temperature for flower

formation is 23C (reviewed in Theron and De Hortogh, 2001).

Mori and Sakanishi (1989) demonstrated that for Agapanthus, flowering occurred

when plants were continuously grown at 10 o or 150C; in contrast, flowering was inhibited

when plants were grown at 200C or higher from September onward. They also found that

flowering could be accelerated by growing plants at a minimum of 20C after the end of

November in combination with a 16-hour photoperiod.

Hartsema (1961) reported that flower formation of Zepi, h)Iinthel, rosea occurred at

13 to 180C and flowering occurred at 220C, with soil moisture being important. No

flowering occurred at 30C. In 7Zeh,1yi n1,the candida inflorescences were formed in

spring and when plants with at least two previously formed inflorescences were subjected

to 100, 15, 23 or 30C in October, earliest flowering occurred on plants exposed to 23C

(Hartsema, 1961).

Vernalization

Vernalization is a period of cold temperature treatments that accelerates flowering

in some plant species. Many biennial (two-year) plants require a temperature below a

critical level for a definite time period before flowering can occur. Plants can either have

an obligate or quantitative vernalization period. Obligate vernalization refers to plants

that require cold temperatures for a period of time in order to flower. Quantitative

vernalization refers to plants that do not require cold temperatures to flower but start their

flowering period earlier under cold temperatures. Additionally, there are plants that do

not respond to cold temperatures (reviewed in Yan and Wallace, 1995).









Flower Initiation Process in Bulbous Plants

Most true bulbs have a sympodial branching system (superposed branches) and at

flower initiation a growing point is formed laterally in the apex. This growing point

develops a certain number of leaves before an inflorescence develops, alternating flower

and leaf formation throughout the entire growth period (Hartsema, 1961). The

flower/inflorescence emergence is delayed compared to leaf emergence thus leaves and

flower/inflorescence above ground have a difference of one generation between them,

and the inflorescence appears to be lateral to leaves (De Hertogh and Le Nard, 1993).

Blaaum (1931) described the general flowering process in true bulbs and according

to his studies bulbs undergo the following stages: I) the meristem is vegetative and

produces leaves; II) the last leaf and new growing point are formed; III) a certain number

of leaves are formed; IV) then the flower/inflorescence meristem is formed; followed by

flower and floral parts formation. More recently, De Munk and Van der Hulst (cited in

Theron and De Hertogh, 2001) have described the flowering process as: Stage I)

vegetative; Stage II) formation of spathe; Stage III) beginning of flower initiation; Stage

IV) flower development and anthesis; Stage V) flower senescence and vegetative growth.

Theron and De Hertogh (2001) showed that Hippeastrum bulbs initiate the

flowering process in the spring, the differentiation period lasts from 18 to 24 months and

anthesis occurs in the spring.

Flower Bulb Cultivation

Field production of bulbous crops occurs where soil and climate have advantageous

characteristics. A large bulb industry developed in the Northwest region of the United

States because soils usually do not freeze enough to damage the bulbs and abundant

rainfalls create favorable growing conditions for them. Mild temperatures and abundant









moisture favor the production of bulbs having a multi-year production cycle. In more

severe climates, such as the Midwest and Northeast, tender bulbs must be removed from

the soil in the fall and stored during the winter (Johson et al., 1995).

Some bulbs grow well in light sandy or gravelly-type soils. However, most bulbs

grow best in loams with high organic matter content. Generally, bulbs do not grow well

in water-logged or heavy clay soils (Johson et al., 1995).

The majority of bulbous crops are planted in the early fall; however, some varieties

are planted during the spring, such as Gladiolus, Gloxinia and Begonia. The bulblets

remain in the ground for 1 to 3 years until they reach harvestable size. Some tender

species require "lifting" or removal from the soil in the fall to avoid freeze damage

(Johson et al., 1995).

The harvesting process is usually done mechanically, using methods similar to

those used in onion production. Bulbs are lifted from the soil and deposited onto a belt-

conveyor that moves them into the harvester, which shakes the bulbs to loosen and

separate the soil. After harvesting, bulbs are sorted, graded, and damaged bulbs are

discarded (Johson et al., 1995).

Flower bulbs present several different growth habits and when cultivated under

growing conditions much different than their native habitat, they may drastically change

their growth and flowering habit. For example, Ornithogalum, which is a deciduous

perennial, becomes an evergreen when grown in the tropics, does not go dormant, blooms

constantly, and does not produce bulbs (Halevy, 1990).

Flower Bulb Forcing

Forcing is defined as the regulation of bulb growth and development under

greenhouse controlled environmental conditions (De Hertogh, 1977). True bulbs, as well









as corms, rhizomes, and tubers can be forced. According to De Hertogh (1977), in order

to succeed using this process a grower must fully understand: a) the origin and

morphology of the species, b) the production and growth cycle of the species, and c) the

influence of various environmental factors on the development of the species.

The complete forcing system has been divided into four distinct phases: I)

production, II) programming, III) greenhouse and IV) marketing. The exact programming

of temperatures and times varies from species to species and might not be applicable to

all cultivars of all species. The complete process should be based on marketing time and

bulbs should finish the process at proper stage of development (De Hertogh, 1974).

The basic bulb production cycle initiates when bulbs are harvested, sorted and

graded (according to their circumference measurements). The process continues with the

storage of bulbs in warm temperatures to fully develop the floral organs. In the fall, the

bulbs are planted, kept moist and under temperatures low enough to promote flowering

and bulbing. In spring, the floral stalk elongates and the plant flowers. Some species

require the removal of flowers, to increase bulb size (De Hertogh, 1977).

Successful forcing of flowering bulbs in a greenhouse is based on seven factors:

temperature (the most important), watering, light, fertilization, ventilation, sanitation, and

pest control. Forcing can be either accelerated or delayed by manipulating these factors

(De Hertogh, 1996).

Hippeastrum bulbs can be forced for either fresh cut flowers or flowering potted

plants. The key factors for forcing Hippeastrum bulbs are: 1) use bulbs larger than 20 cm

in circumference (6 cm in diameter), 2) remove the bottom half of the roots, 3) cure the

bulbs for 10 days at 170 to 230C, 4) store the bulbs at 90 to 130C for at least 8 weeks









before planting, 5) package bulbs to avoid roots drying out if transported, 6) plant bulbs

in well-drained planting medium, 7) use bottom heat of 22C in the greenhouse (reviewed

in Theron and De Hertogh, 2001).

Tropical Bulbs and Amaryllidaceae

Most tropical bulbs, and in particular those used in this study, belong to the family

Amaryllidaceae, which is a very diverse family with species on almost all continents and

under various climatic conditions (Meerow and Snijman, 1998).

Amaryllidaceae classification includes the following:

Kingdom: Plantae

Subkingdom: Tracheobionta (Vascular plants)

Superdivision: Spermatophyta (Seed plants)

Division: Magnoliophypa (Flowering plants)

Class: Liliopsida (Monocodiledoneous)

Subclass: Liliidae

Order: Liliales / Amaryllidales / Asparagales

Family: Amaryllidaceae

According to Watson and Dallwitz (1992) the family Amaryllidaceae comprises

approximately 60 genera and 800 species. Some of the important horticultural crops are

Amaryllis, Clivia, Crinum, Eucharis, Haemanthus, Hippeastrum, Hymenocallis, Lycoris,

Narcissus, Nerine, Scadoxus and Ze7p,h)y 1i1he,.

The two most recent formal classifications of the Amaryllidaceae family are those

of Muller-Doblies and Muller-Doblies (1996) and Meerow and Snijiman (1998).

Hyacinthaceae, is also considered a related family, however Agavaceae, Hypoxidaceae,









Haemodoraceae or Alstroemeriaceae with which Amaryllidaceae is sometimes

associated, are not considered related families.

Plants in the Amaryllidaceae family have distinct habits, however most of them are

perennial, bulbaceous and have contractile roots. Leaves are mostly deciduous and

simple, with entire lamina, entire margins and parallel venation. Flowers may be solitary

or produced on different types of inflorescences. Bulbous plants in this family have

thickened underground storage organs which enable them to survive unfavorable

environmental conditions and may also function as propagative units (reviewed by

Theron and De Hertogh, 2001).

Halevy (1990) demonstrated that when flower bulbs are produced under growing

conditions that are dissimilar to their indigenous environments, their growth habit can be

altered. Therefore, it is important to understand the effects of temperature, light, nutrition,

growth regulators, and other environmental factors on bulb growth, development and the

flowering process, to significantly expand the horticultural usage of the Amaryllidaceae

family (Theron and De Hertogh, 2001).

Hippeastrum spp.

The name Hippeastrum comes from the Greek hippeus, meaning knight and astron

meaning star. Hippeastrum is an important genus of the Amaryllidaceae family which

comprises about 70 species, such as H. argentinum, H. aulicum, H. barbatum, H.

correiense, H. elegans, H. evansiae, H. leopoldii, H. miniatum, H. morelianum, H.

pardinum, H. psittacinum, H. puniceum, H. reginae, H. reticulatum, H. rutilum, H.

stylosum, H. vittatum, and H. reticulatum (Rees, 1985). There are more than 300 cultivars

and most of the horticulturally important ones were bred by Ludwig, Warmenhove and









van Meeuven in the Netherlands and by HADECO (Barnhoorn) in South Africa (Read,

2004).

Hippeastrum can be considered a tropical plant by origin as it is indigenous to

Central and South America, being centered in Brazil and Peru, and distributed from

Mexico to Argentina (De Hertogh and Le Nard, 1993). Hippeastrum is commercially

known as Amaryllis, however, the true Amaryllis (A. belladonna) originated in South

Africa and it is not widely used. There are several differences between them such as the

presence of a solid scape and the absence of scales between the filaments in A.

belladonna (Goldblatt, 1984).

Hippeastrum spp has large and showy flowers with many bright colors (red, pink,

orange, white or bi-colored) consisting of several flower types: trumpet-flowered,

belladonna types, reginae types, leopoldii types, miniatures, doubles, and orchid-

flowered. They are grown mainly as potted plants or as cut flowers, but they can also be

grown in the landscape in subtropical and tropical areas (Schulz, 1954).

The bulb has a sympodial branching system, which means that the terminal bud

dies or ends in an inflorescence, and growth of sympodial shoots continues from lateral

buds. At flowering initiation, a lateral growing point is formed on the side of the apex. It

develops in a sequence of four leaves and an inflorescence. Bulblets are initiated in the

axils of senescing bulb scales in the outer parts of the bulb and they produce nine leaves

before initiating the first inflorescence (reviewed in Theron and De Hertogh, 2001).

Scadoxus multiflorus Blood Lily

The genus Scadoxus contains 9 species: S. cinnabarinus, S. cyrtanthiflorus, S.

longifolius, S. membranaceus, S. multiflorus, S. nutans, S. pole-evansii, S. pseudocalus,

and S. puniceus (Friis and Nordal, 1976). Scadoxus is closely related to Haemanthus;









they were treated as a single genus at one time but were divided by Friis and Nordal

(1976). The major differences between Scadoxus and Haemanthus are related to their

geophytic organs, growth habits, and number and shape of their leaves (Snijman, 1984).

Scadoxus multiflorus is one of the most horticulturally relevant species in the

genus, its name derived from doxus meaning glory or splendor, and multiflorus referring

to many flowers (Jackson, 1990). Scadoxus multiflorus is endemic to southern and central

Africa and was established by Rafinesque (1836). This species includes three subspecies:

katherinae, longitubus and multiflorus, and the major differences between the subspecies

are height of the plants, length of the perianth tubes and the extent of the perianth

segments (Du Plessis and Duncan, 1989).

Agapanthus africanus African Lily

The genus Agapanthus was established by L'Heritier (1788). The name

Agapanthus is derived from the Greek agape that means love and anthos that means

flower (Snoeijer, 2004). It was placed in the Liliaceae family, later moved to the

Amaryllidaceae family. Agapanthus is endemic to southern Africa and the first species

collected Agapanthus africanus was described in 1679 by the name Hyacinthus

africanus tuberosus, which is sometimes still referred to as Agapanthus umbellatus

(Leighton, 1965).

Agapanthus is a variable genus, all species are rhizomatous with similar

appearances. Botanists consider it difficult to classify them into distinct species. Plessis

and Duncan (1989) identified about 10 species indigenous to Southern Africa.

Agapanthus africanus is a summer-flowering evergreen species, with its perennial

geophytic organ a rhizomatous rootstock with contractile roots. Flowers are formed in an

umbel inflorescence on a leafless scape. Mori and Sakanishi (1989) observed that A.









africanus meristem was vegetative in October, flower initiation began in November and

flower differentiation occurred in December.

Habranthus robustus and Zephyranthes spp. Rain Lilies

Habranthus robustus was established by Herbert and Loddiges (1831), and

Ze7',1)1u/inwhe, spp. was established by Herbert (1821). The name Habranthus comes from

the Greek habros, meaning graceful and anthos meaning flower, and 7Zeqhy/tliin comes

from the Greek, zephyros meaning "the west wind" and anthos, meaning flower. These

two groups of summer-flowering small bulbs of the Amaryllidaceae family are native to

the southeastern United States, Central and South America (Hume, 1935; Traub, 1958 b).

The genus Habranthus contains about 40 species, and among the horticulturally

important are H. tubispathus and H. robustus, which is native to Brazil (Read, 2004). The

Ze7,1h)1 l-ihe, genus contains about 60 species and the most horticulturally important are

Z. candida Herb and Z. grandiflora Lindl, but other commercially grown species include:

Z andersonii Baker, Z atamasco (L.) Herb, Z drummondii, Z primulina T.M. Howard

& S. Ogden, Z rosea Lindl, and Z. citrina Baker. There are several named cultivars

derived from interspecific crosses (Van Scheepen, 1991).

Habranthus and Ze7,1'l) ,iahie geophytic organs are perennial bulbs covered with a

dark tunica, contractile roots, deciduous leaves with sheathing basis and linear blades,

and isolated flowers (Theron and De Hertogh, 2001).

Habranthus robustus flowers are very similar to those of Zep,1y )l, ihe/ and

Habranthus have at times been included in the genus Ze7,1p) iirihe because of that. Both

are commonly called "rain lilies" because of their tendency to bloom after rain periods

(Fellers, 1996).









Habranthus robustus is often called Ze7,1'/i-iihi/e robusta incorrectly, which adds

to the confusion between it and Ze7,lo,1i ainhe grandiflora. The very subtle differences

between genera are based on spathe characters, position of flowers, symmetry of corolla,

insertion of anther filaments, number and length of filaments, and number of seeds per

locule in capsule (Femandez-Alonso & Groenendijk, 2004).

Ze7,1)1,o iihe' flowers point straight up and have equal lengths stamens, while

Habranthus flowers point upward but at an angle and have stamens of different lengths;

these characteristics are commonly used to separate the two genera. Additionally,

Habranthus flowers tend to have zygomorphic flowers (bilaterally symmetrical), while

Ze7,1'l)I -ihe, are actinomorphic flowers (radially symmetrical). The flowers of

Habranthus are clearly distinguished by filaments of 4 different lengths and always

longer than the perianth tube. Ze7,1yhl)iinhe, flowers are declinate and distinguished by

filaments of two very similar lengths (Femandez-Alonso & Groenendijk, 2004).

Ze7', l)I/1ahe, and Habranthus can also be distinguished by the shape of their seed.

Ze7,1)*/o1iiw/ seeds tend to be more D-shaped or wedge shaped while those of

Habranthus are more openly obliquely winged. These two genera are also distinguished

phylogentically as, Ze7,1oiji ,nhe, apparently have 2-3 different origins according to

nrDNA spacer sequences (Meerow et al, 2000).

The differences between Habranthus and Ze7,1') ,iirhe' are not readily apparent to

most growers and consumers but the two groups of bulbs seem to have different

flowering periods and re-blooming characteristics. The results of this study will

contribute to a better understanding of these species and their cultural differences. This






30


study will also help growers determine which species grow best at different times of the

year and maximize seasonal sales and facilitate their commercialization.














CHAPTER 3
BULB MORPHOLOGY

Geophytes were defined by Raunkier (1934) as plant species with specialized

underground storage organs that accumulate food reserves, nutrients and moisture for

seasonal growth and development. They are usually collectively called "flower bulbs"

and can be separated into four groups: true bulbs (tunicate and non-tunicate), corms,

tubers and rhizomes. Although morphologically different, all types of geophytes perform

the same basic function, storage of photosynthates.

Most geophytes have a shortened stem, a basal plate, one or more apical meristems,

enclosed flower buds, adventitious root initials, several layers of fleshy scales (modified

leaves), and a protective tunic. The size of storage organs vary tremendously among

species (Proches et al., 2006), as well as their leaf arrangements and flower formation.

Some genera, including Tulipa and Narcissus (Caldwell and Wallace, 1955) have been

extensively examined. However, only limited information is available in the literature

concerning morphology of other tropical bulbs.

This portion of the present study was designed to compare and contrast the

morphology and flower formation of Hippeastrum hybridum, Scadoxus multiflorus and

Agapanthus africanus, and compare them to Habranthus robustus and Zelphl)1,uie,% spp,

which were species used in the subsequent phases of this study.

Comparative Study

Materials and Methods: The five species of tropical bulbous plants listed on table

3-1 were grown under same conditions, dissected by freehand sections, stained with









Safranin, and examined during mature and immature stages of development. Digital

images and drawings were produced to determine similarities and differences in

morphology and floral development of the five species examined.

Table 3-1 Tropical geophytes used in the morphological studies:
Species Common Name Family Source
Hippeastrum hybridum. Amaryllis Amaryllidaceae Agristarts, Inc.
Scadoxus multiflorus Blood Lily Amaryllidaceae Agristarts, Inc.
Agapanthus africanus Agapanthus Alliaceae Agristarts, Inc.
Habranthus robustus Rain Lily Amaryllidaceae UF stock
Zep,1ij ,i uhei spp. Rain Lily Amaryllidaceae UF stock


Seventy two plugs ofHippeastrum hybridum, Scadoxus multiflorus and

Agapanthus africanus were obtained from Agristarts Inc. in Apopka, Florida during the

first week of March 2004. Habranthus robustus and Zep,1i, lilrn/i hybrids bulbs were

obtained from University of Florida stock where they had been grown in ground beds for

a year prior to the beginning of this study. The 7Zephyiin,1he hybrids used in this study

were: Z. 'Paul Niemi', Z. 'Jo Ann's Trial' and Z. 'Fadjar's Pink', which were hybridized

by Fadjar Marta in Jakarta, Indonesia.

All geophyte plugs and Habranthus and Zep,1lyii ,ulni bulbs were transplanted into

15 cm plastic pots using sphagnum peat based Fafard No. 2 soiless growing medium

(Agawam, MA) consisting of 70% Canadian sphagnum peat, 10% perlite and 20%

vermiculite. Plants were placed in a greenhouse (which provided 11% shade), natural

photoperiod and a temperature range of 31/24C (day/night). From planting through

establishment all pots were irrigated every other day with 250ml of water, except

Hippeastrum which was watered daily with same amount of water. After establishment

(first week of April), plants were fertigated twice a week with 250ml of water containing









Peters Professional 'Florida Special' water soluble fertilizer 20N-4.7P-16.6K (Scotts Co.,

Marysville, OH) with N at 150 mg L-1. Plants were grown in the greenhouse for ten

months.

From June to November 2005, a series of freehand sections of the four species of

bulbs and Agapanthus rhizomes were made. The sections were mounted in glycerin and

stained with Safranin for 24 hours to facilitate the observation of specific tissue

components. Sections were examined with a Wild M5 dissecting scope and light

microscopy images were obtained with a Zeiss Tessovar with an Rts Contax camera

attachment.

Freehand sections of the meristematic regions of the five species studied were

mounted in glycerin (but not stained) and light microscopy images were obtained with a

Nikon Labophot-2 microscope and a Nikon E4500 digital camera attached to it.

Drawings, based on observations of numerous freehand sections, were made to

demonstrate the general anatomy and morphology of each of the five species examined.

Results and Discussion

Bulb type and size: Observations and measurements made in this study revealed

that Zep7,1/ltwihe/ and Habranthus bulbs were similar in shape but that Zpyl/iiiihe

bulbs were smaller than Habranthus bulbs. Mature Z7,p/tiiiinew\ bulbs were

approximately 6 cm tall with a diameter of about 6 cm, while mature Habranthus bulbs

were approximately 8 cm tall with a diameter of about 8 cm. Both bulb genera were

covered with a fine, papery dark brown tunica. Hippeastrum bulbs were covered with a

fine, papery dark brown tunica and were similar to Z7,p1iyin1whe and Habranthus in

shape but were larger, as mature bulbs were approximately 12 cm tall with a diameter of

about 12 cm (Table 3-2 and Figure 3-1). Zephyranthes, Habranthus and Hippeastrum









form true tunicate bulbs, with concentric layers of modified leaves called scales (Figures

3-2, 3-3 and 3-4).

Scadoxus geophytic organs were true bulbs formed by concentric layers of scales

(approximately 6 cm tall when mature) with a thick (4 cm tall) rhizomatous structure at

their base and covered by a fine, papery dark brown tunica (Figure 3-5). Scales of

Scadoxus were thicker (0.5 cm) than scales of Zephyranthes, Habranthus and

Hippeastrum which averaged about 0.2 cm in thickness.

Agapanthus geophytic organs were rhizomes and did not have layers, scales or a

tunica as did the other four previous studied species (Figure 3-6). A mature Agapanthus

rhizome was approximately 20 cm long with a diameter of approximately 10 cm (Table

3-2).

Table 3-2 Geophyte size and characteristics of plants examined
Genera Geophytic Height Diameter Scales Tunica

organ (cm) (cm) present present

Hippeastrum True bulb 12 12 Yes Yes

Habranthus True bulb 8 8 Yes Yes

Zepi7,yiu)1, True bulb 6 6 Yes Yes

Scadoxus Bulb/rhizome 10 12 Yes Yes

Agapanthus Rhizome 10 No No































figure 3-i: rrom let to rignt Lepnyrantnes, naorantnus, nippeastrum, 3caaoxus DUIDs,
and immature Agapanthus rhizome. Bar = 10 cm.


(A) (B)
Figure 3-2: Sections of Zepi, hy ,uthe bulbs. Longitudinal section (A) and
photomicrograph of a cross section (B). Bar =1 cm.






















(A) (B)
Figure 3-3: Sections of Habranthus bulbs. Longitudinal section (A) and photomicrograph
of a cross section (B). Bar = 1 cm.













(A) (B)
Figure 3-4: Sections of Hippeastrum bulbs. Longitudinal section (A) and
photomicrograph of a cross section (B). Bar = 1 cm.














(A) (B)
Figure 3-5: Sections of Scadoxus bulbs Longitudinal section (A) and photomicrograph of
a cross section (B). Bar = 1 cm.


























(A) (B)
Figure 3-6: Longitudinal sections of an Agapanthus rhizome. Bar = 1 cm.

Leaf arrangement and morphology

Hippeastrum, Habranthus and Zep,1,Iliu/1ihe bulbs had similar leaf arrangements

and leaf morphologies. Leaves were perennial, basal, simple, linear, glabrous, with entire

margins and parallel venation. They emerged from the center of the bulb, with leaves

two-ranked as the blade of each new leaf emerged 1800 from the previous leaf, thus older

leaves were always on the outside and younger leaves on the inside of the leaf cluster

(Figure 3-7, 3-8 and 3-9). Each leaf was composed of a photosynthetic leaf blade and a

non-photosynthetic storage leaf base (scale), which thickened during the growth cycle

forming true bulbs.

Jones and Emsweller (1936) stated that each new leaf ofAllium cepa developed on

the side of the apical meristem opposite to the preceding blade by an upward growth of

tissue surrounding the apical meristem. We observed that Hippeastrum, Habranthus and

7Zepi2,/),1ihe, leaf production occurred in a similar process. According to Mann (1960) A.

cepa bulbs contain leaves that are morphologically distinct from each other, and









Kamenetsky (1994) classified them according to their function: protective, storage and

assimilation.

Black et al. (1990) described Hippeastrum leaf blades as being up to 60 cm long

and up to 5 cm wide. We observed that Habranthus leaf blades were 10 to 20 cm long

and 1.5 to 2 cm wide; while Zepi7,yi,)i/he, blades were 10 to 20 cm long and 1 cm wide.

As part of this study, a determination of number of leaves produced weekly for a period

of one year of Habranthus and Zei7y,1iu)i/we, bulbs was made (Figures 3-41 and 3-42).

Agapanthus rhizomes have simple, linear, perennial glabrous leaves with entire

margins and parallel venation. Leaf blades were up to 50 cm long and approximately 4

cm wide. Leaf arrangement was alternate but we observed that the leaves did not emerge

from the center as they did in Hippeastrum, Habranthus and Zephyranthes, but from the

lateral sides of the rhizome (one at a time). Thus, the older leaves were external to inner

new leaves. Another distinction between Agapanthus and Hippeastrum, Habranthus and

Zep7,yiirwhe,1 bulbs was that leaf bases did not thicken and form scales, but remained thin

and papery (Figure 3-10 C).

Scadoxus leaves were shorter, wider and thinner than leaves of Hippeastrum,

Habranthus, 7Zephyihtu1he, and Agapanthus, and they had a distinct oblong shape,

prominent mid-ribs, undulating leaf margins, and leaf blades were 30 to 40 cm long and 9

to 13 cm wide. Scadoxus leaf arrangement was distinct from the previous four species.

The first two leaves emerged singularly and were distinctly different from subsequent

leaves. After the first two leaves senesced, a cluster of leaves arose from the center of the

bulb all at once. They were held together by a structure called a pseudo-stem (a false

stem formed by the sheathing and overlapping of the leaf bases Figure3-11 C).










Species Leaf blade length (cm) Leaf blade width (cm)

Hippeastrum Up to 60 5

Habranthus 10 to 20 1.5 to 2

Zep7iyumi/e, 10 to 20 1

Scadoxus 30 to 40 9 to 13

Agapanthus Up to 50 4


(A)' (B(C
Figure 3-7: Hippeastrum leaf arrangement. From top (A), side (B) and individual leaf
with continuous leaf base / scale (C). Bar = 1 cm.












(A) (B)
Figure 3-8: Habranthus leaf arrangement. From top (A) and side (B). Bar = 1 cm.


Table 3-3


I.eaf si Ze of nlants examined


















(A) (B)
Figure 3-9: Ze7,1phi)/1,the leaf arrangement. From top (A) and side (B). Bar = 1 cm.












(A) (B) (C)
Figure 3-10: Agapanthus leaf arrangement. Overlapping of leaf bases (A), new leaves
emerging from lateral side of the rhizome (B), and individual leaf with
continuous papery leaf base (C). Bar = 1 cm.

.A/ -









(A) (B) (C)
Figure 3-11: Scadoxus leaf arrangement. First emerged leaf (A), photomicrograph of a
longitudinal section showing leaf sheathing in a pseudo-stem (B), and a group
of mature leaves in a pseudo-stem (C). Bar = 1 cm.









Floral initiation

Free hand cross sections of the bulbs used in this study revealed that floral bud

formation was similar in Hippeastrum, Habranthus and 7Zeh1yii unhe, bulbs (Figures 3-

12, 3-13 and 3-14). Flower buds were formed centrally in the apical meristem, and were

initiated during the bulb's vegetative growth but did not emerge until the bulb reached a

critical size. The elongation of flowering scapes occurred after the formation of 3 to 4

leaves on these species. We observed that flower buds were produced alternatively from

each side of the meristem and formed a cross line in cross section (Figure 3-15). Since

the apical meristem was constantly active (except during dormant periods) the older

flower buds were located on the outer part of the bulb, identical to the leaf development

pattern of these three species (Figure 3-16).

The average number of leaves initiated prior to inflorescence initiation depends

upon the species in Amaryllidaceae bulbs. Hartsema and Leupen (1942) found that 11

leaves were initiated before floral initiation in Amaryllis belladonna and 8 in Nerine

bowdenii; while there were 4 in Hippeastrum hybridum (Blaauw, 1931), 15 in Scadoxus

multiflorus (Peters, 1971), 7 to 8 in Leucojun aestivum L. (Luyten and Van Waveren,

1938), 5 in Lycoris radiata and 10 in Lycoris squamigera (Mori and Sakanishi, 1977).

We observed that there were 4 leaves formed between each inflorescence in both

Habranthus and Zep7,jhy)i/,nhi bulbs (Table 3-4).

In Allium cepa, which is similar to Hippeastrum, Habranthus and Zephyranthes,

the first stages in the development of the leaf and inflorescence primordia in the

meristematic region are similar and an inflorescence bract is indistinguishable from a leaf

blade, as both are protected by an involucre and a series of bracts (Jones and Emsweller,

1936).









Hippeastrum flowers are produced in groups of 2 to 6 in an umbel inflorescence

and are attached to a hollow scape (around 50 cm tall) that can appear singly or more

than one at the same time. A mature bulb produces about 12 leaves and 3 to 4 scapes per

season. Inflorescences in Habranthus and 7Zep ,yihihe1 \ possess only one flower in most

cases (rarely 2 flowers) attached to a hollow scape. We observed that a mature bulb can

produce up to 4 scapes at the same time (figure 3-17). Both Habranthus and

Z.7',1/)1r/iahe, bulbs can flower several times during one season, their flowers last for 2 to

3 days and leaves may or not be present during flowering. Throughout this study it was

observed that it takes four days for a Habranthus flower to transition from visible flower

bud to an open flower (Figure 3-18).

In Scadoxus bulbs, flower buds and pseudo-stems are formed side by side at the

apical meristem (Figure 3-19), one of each per season. The inflorescence scape which is a

spherical umbel (25 cm in diameter) consisting of up to 200 flowers, emerges first and

then the foliage held by a solitary pseudostem emerges (Figure 3-20). The inflorescence

lasts for 1 to 2 weeks, while the pseudo-stem lasts for a few months. Their flowering

season is in late summer to early autumn.

We observed that Agapanthus rhizomes did not produce inflorescence and leaves in

the same region of the basal plate. Inflorescences were formed in the apical meristem

located in the central part of the basal plate while leaves were developed from apical

meristems distributed along the lateral sides of the basal plate (Figure 3-21A).

Agapanthus inflorescences are umbels with 8 to 20 flowers held by stiff erect scapes that

arise from the center of the rhizome (Figure 3-21B). Inflorescences last for 1 week and

flowers appear during summer.









Table 3-4


-Relationship between leaf and flower formation, and number of flowers per
iniflorescence of plants examined


Species Leaves between inflorescences Number of flowers per inflorescence

Hippeastrum 4 2 to 6

Habranthus 4 1 to 2

Zep7,ji ihe,' 4 1 to 2

Scadoxus 15 Up to 200

Agapanthus 8 to 20


-
(A) (B)MKP t
Figure 3-12: Longitudinal sections of a Hippeastrum bulb. Free-hand section (A) and
photomicrograph (B). Bar = 1 cm.


(A) (B)
Figure 3-13: Sections of Habrantus bulbs. Inner part with outer scales removed (A) and
photomicrograph of a longitudinal section (B). Bar = 1 cm.






















(A) (B)
Figure 3-14: Sections of Zep,1/it wllhe, bulbs. Inner part with outer scales removed (A)
and photomicrograph of a longitudinal section (B). Bar = 1 cm.


Figure3-15: Cross section of a Habranthus bulb (stained with Safranin), showing the
alignment of flower buds. Bar = 1 cm.































Figure 3-16: Longitudinal section of a Habranthus bulb (stained with Safranin), showing
flower buds. Bar = 1 cm.


Figure 3-17 Habranthus bulb with three flower stalks. Bar = 10 cm.




















(A, (13)












(C) (D)
Figure 3-18 Development of a Habranthus flower. Flower bud (circled) emerged from
soil on day 1 (A), flower stalk elongated (circled) on day 2 (B), flower stalk
(circled) elongated on day 3 (C) and open flower on day 4 (D). Bar = 5 cm.


Figure 3-19: Sections of Scadoxus bulbs. Inner part with outer scales removed (A) and
photomicrograph of a longitudinal section (B). Bar = 1 cm.






























Figure 3-20: Flower st;
1 cm.


emerged and pseudo-stem arising from a Scadoxus bulb. Bar


A(A)MZP (B). .
Figure 3-21: Sections ofAgapanthus rhizomes. Photomicrograph of an in longitudinal
section (A) and flower stalk emerging from center of an Agapanthus rhizome
(B). Bar = 1 cm.









Meristematic region

The apical meristem consists of a group of undifferentiated cells located at the apex

of a shoot or a root that continuously produces new cells. The apical meristem was first

observed in 1759 by Caspar Wolff, who recognized it as being the center of organ and

cell formation (Tooke and Battey, 2003).

There are four types of meristems: single apical cell or initial (that can be lenticular

or tetrahedral), cluster of apical cells, zoned apex and tunica corpus type. The first two

types are characteristic of ferns and other lower plants, the third type is characteristic of

gymnosperms, and the fourth type occurs in angiosperms. The five species of tropical

bulbs studied have the tunica-corpus type of shoot meristem (Dengler, 2002).

Studies of meristems reflect the technology of the time period in which they were

conducted. During the 19th and 20th centuries, observational analysis were made on

sections of living and preserved tissue, from 1940 to 1970, experimental manipulations

were made with microsurgery, radioisotopes, labeling and chimeric analysis. Since 1970,

genetic and molecular analyses have been employed to gain information on plant

meristems (Tooke and Battey, 2003).

Tulip apices were described by Sass (1944) as a tunica-corpus type, during the

vegetative state and flower differentiation. The vegetative apex of the principal axis is

deeply buried in the bulb, just above the basal plate, and it is 2 to 3 mm long and 1 to 2

mm wide. The apex is a short dome 100 to 125 itmm high and 300 to 375 itmm broad

and approximately semicircular in longitudinal section. The apical meristem of Allium

cepa has also been described as a low, circular, dome-shaped mass of cells (Hoffman,

1933).









In tulips, the tunica consists of a single layer of cells, followed by several layers of

the corpus. The leaf primordia results from division of cells in the first layers of the

corpus and rapidly involve zone three or four cells deep. According to Rees (1972),

different plants can have a tunic with more than one layer of cells. That is the case in Iris,

which posses a tunica of two layers of cells, and Lilium candidum, which has a tunica

formed by three layers of cells. This study did not identify the number of cell layers in the

tunica of each species but clarified the cellular layout of the apical region in all five

species studied.

Hippeastrum, Habranthus, Ze7,1pyt1uihe, and Scadoxus bulbs had a similar

arrangement and their apical meristematic regions were located in the basal plate, as

described for Allium cepa (Mason, 1979). The apical meristem and youngest leaf

primordia were surrounded by scales (Figures 3-22 to 3-29). Blaaum (1931) found that

the first morphological sign of inflorescence initiation in Hippeastrum bulbs was a slight

increase in the size of the apical meristem.

Agapanthus rhizomes were distinctly different that Hippeastrum, Habranthus,

Ze7,1pl)I rihe, and Scadoxus, as they had a centrally located apical meristem responsible

for flower formation and several lateral meristems located on the sides of the rhizome,

which were responsible for leaf production (Figures 3-30 and 3-31).







50









Bulb Scale
(Swollen leaf base)

Developing Leaf

Developing Leaf

Leaf primordium
Leaf primordium
Meristem
Flower bud

Vascular channel
Basal plate




Figure 3-22: Photomicrograph of the meristematic region of a Hippeastrum bulb. Bar = 1
cm.








Developing leaf

Unmerged leaf


Leaf primordium

Leaf primordium
Meristem

Vascular
channel

Basal plate





Figure 3-23: Photomicrograph of the meristematic region of a Hippeastrum bulb. Bar =1
cm.







51








Bulb scale
Swollen leaf base

Unmerged leaf

Develo ing leaf



Leaf primordium
Meristem

Vascular channel

Basal plate




Figure 3-24: Photomicrograph of the meristematic region of a Habranthus bulb. Bar = 1
cm.








Developing leaf

Unmerged leaf


Leaf primordium

Leaf primordium
Meristem

Vascular
channel

Basal plate





Figure 3-25: Photomicrograph of the meristematic region of a Habranthus bulb. Bar = 1
cm.







52






Flower bud
Developing leaf t


Flower bud f
Leaf primordium

Meristem

Vascular channel

Basal plate .f "







Figure 3-26: Photomicrograph of the meristematic region of a Ze7,hi ihie'\ bulb. Bar
1 cm.






Developing leaf

Developing leaf
Flower bud
Leaf orimordium
Leaf primordium
Meristem



Vascular channel
Basal plate




Figure 3-27: Photomicrograph of the meristematic region of a Ze7,h)i whie' bulb. Bar
1 cm.


















Developing leaf

Developing leaf


Leaf primordium

Leaf primordium

Meristem

Basal plate

Vascular channel


Figure 3-28: Photomicrograph of the meristematic region of a Scadoxus bulb. Bar = 1
cm.










Develo in leaf

Develop in leaf

Leaf orimordium
Leaf primordium
Meristem

Vascular channel '

Basal plate






Figure 3-29: Photomicrograph of the meristematic region of a Scadoxus bulb. Bar = 1
cm.







54










Emer ed flower stalk

Developing leaf


Leaf primordium .
TV I

Lateral meristem .
Leaf primordium
Lateral meristem .

Leaf rimordium t',, .i
Lateral meristem
Basal plate /sa
Vascular channel

Figure 3-30: Photomicrograph of the meristematic region of an Agapanthus rhizome. Bar
= cm.









Develooinc leaf

Developing leafr



Leaf primorJLuIIn
Leaf primorJLuIIn
Lateral menstemn

Vascular channel
Basal Plate




Figure 3-31: Photomicrograph of the meristematic region of an Agapanthus rhizome. Bar
= cm.










General anatomical and morphological arrangement


Uneience.ld c.l --


Basal plate


~Co..frJ~i. i


Figure 3-32: Longitudinal section of a Hippeastrum bulb showing its general anatomy
and morphology. Bar = 1cm.


;I.,r / **.,
.; ..'o. ,
'"
-/


,/

/


Basal plate / I _

Figure 3-33: Longitudinal section of a Hippeastrum bulb showing its general anatomy
and morphology. Bar = 1cm.

























Figure 3-34: Longitudinal section of a Ze7',1ji,/ille, bulb showing its general anatomy
and morphology. Bar = 1cm.

Bulb scale
Unemerged l.c -

Flower bud


( / I t
V" ; /, *'- V "


~~K


Basal plate


/ 'O,#, 4t-zR. --


Figure 3-35: Longitudinal section of a Ze7,1' yiilhe bulb. Bar = 1cm.


























Basal plate
/^-;'^-- /;,,-._. -


(A) (B) I
Figure 3-36: Sections of Habranthus bulbs. Longitudinal (A) and transversal (B)
sections, showing the reduced basal plate with leaves/scales between flower
buds (A) and a series of leaves/scales (B). Bar = Icm.







Psc udo stem

Bulb scale


Bai plate









Figure 3-37: Longitudinal section of a Scadoxus bulb showing its general anatomy and
morphology. Bar = Icm.











Unemerged leaves

Pseudo stem

-j Bulb scale

Basal plate

















Figure 3-38: Longitudinal section of a Scadoxus bulb showing its general anatomy and
morphology. Bar= Icm.











Emerged leaf

SUnemerged leaf


S- Basal plate


/- Rhizome
t//


Figure 3-39: Longitudinal section of an Agapanthus rhizome showing its general
anatomy and morphology. Bar = 1cm.


S\- \ i / Emerged inflorescence

E. Emerged leaf


Unemerged leaf


SiC j Basal plate





J .kI


Figure 3-40: Longitudinal section of an Agapanthus rhizome showing its general
anatomy and morphology. Bar = 1 cm.









Experiment 1

The storage organs of geophytes are able to supply food reserve for rapid leaf

growth at the beginning of their growth and reproductive cycle, after cold winters and/or

dry seasons (Rees, 1972). In bulbs with hysteranthous leaves (flowers and leaves develop

in separate seasons), an accumulation of storage materials is a prerequisite for flowering

(Burtt, 1970). In bulbs with synanthous leaves, which are present when plants flower, the

storage materials provide for early growth of photosynthetic organs and allocation of

resources for flowering originate from the storage organ and from existing leaves (Dafni

et al., 1981).

Phenological differences can be summarized into two groups based on the speed of

leaf development and duration of photosynthetic period. The rapid route is a

characteristic of bulbs that produce leaves quickly in order to compensate for a short

photosynthetic period or when there is a limiting factor, such as moisture (most

common), light or temperature. The slow route is characteristic of bulbs that produce

leaves slowly, under long photosynthetic periods or when moisture, light and temperature

are not limiting factors (Dafni et al., 1981).

Of the five species used in this morphological study, Habranthus and Zep,10) ra///th

were the least investigated by other researchers, and since these two genera were used in

the subsequent phases of this study an experiment was designed to understand their leaf

and flower production. Habranthus and Ze7,1yi0t1,whe\ have perennial storage organs and

synanthous leaves. According to Dafni et al. (1981) this type of bulb accumulates more

reserves than required for a growth cycle which includes completion of flowering and

seed production even if the net production (by existing leaves) is insufficient. Therefore

relatively slight differences are expected from year to year in the abundance of flowering.









The following experiment was performed to compare leaf and flower production of

Habranthus and Z7,,1)I-iil/w\ during a period of one year.

Materials and Methods: On the first week of January 2005, 12 Habranthus

robustus bulbs and 12 7Zepihyi-inlll bulbs were planted in 15 cm plastic pots, one bulb

per pot, using a sphagnum peat based Fafard No. 2 soiless growing medium (Agawam,

MA) consisting of 70% Canadian sphagnum peat, 10% perlite and 20% vermiculite.

Plants were placed in a greenhouse with 11% shade, natural photoperiod and a

temperature range of 31/24C (day/night). Plants received 250ml of water and were

fertilized at each irrigation with Peters Professional 'Florida Special' water soluble

fertilizer 20N-4.7P-16.6K (Scotts Co., Marysville, OH) with N at 150 mg.L-1. Bulbs used

in this experiment were obtained from the University of Florida stock and had been

grown in ground beds for a year prior the beginning of the experiment.

This experiment was performed from January to December 2005. Data were

collected twice a month during the entire year and the number of leaves and flowers

present at the moment of data collection were recorded. Data was analyzed by taking the

mean number of leaves and flowers per total of bulbs at each observation, using the SAS

statistical package version 8.02 (Cary, NC). Results demonstrate the performance of leaf

and flower production of these two species during 2005.

Results: Habranthus bulbs demonstrated a predictable performance of leaf

production with more leaves at the beginning and end of the year when not flowering and

a gradual reduction of leaf emergence as flowering season occurred (Figure 3-41).

Ze7,h/,l)-1ihe also produced more leaves in the beginning and end of the year, but their

flowering season did not coincide with a period of reduced leaf production. In April









Zep,1 l irihe,' bulbs produced the least number of leaves followed by a gradual increase

until August when a slight drop occurred and the flowering season started. Production of

leaves increased again until October, when another reduction occurred followed by an

increase of leaf emergence until the end of the year (Figure 3-42).

Comparing both figures (3-41 and 3-42) makes it clear that these two species

posses distinct patterns of leaf production. Habranthus bulbs had fewer leaves emergent

when flowering, while leaves of Z7,y'l)1/i,/he, bulbs continued to emerge during

flowering. Overall, 7Zep2,,j1oinihe bulbs had a greater quantity of leaves during the entire

year but with oscillations, while Habranthus bulbs demonstrated a more uniform progress

with a gradual decrease and increase in leaf correlated with the flowering period. This

information is valuable for the landscape use of these two species as they generate

distinct appearances in a garden; 7Zep2ylo)inhe, have copious foliage in combination with

their flowers and Habranthus have few leaves when flowering.








63







* LEAVES 0 FLOWERS


2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 50 52 54
Week

Figure 3-41: Number of leaves and flowers produced by Habranthus bulbs in 2005






SILEAVES *FLOWERS


2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 50 52 54
Week

Figure 3-42: Number of leaves and flowers produced by Zeph7,/1i/the,% bulbs in 2005














CHAPTER 4
ENVIRONMENTAL EFFECTS FERTIGATION FREQUENCY AND FERTILIZER
RATES ON FLOWERING IN Habranthus robustus AND Zp,10 iyiaiwhe spp.

The primary goal of a flower bulb grower is to produce true-to-type, essentially

disease- and insect-free plants that flower successfully. Proper water management and

fertilization are among the critical factors to accomplish that goal.

Most all ornamental bulbs along with onion and garlic (related vegetable crops) are

sensitive to water deficit. When the soil is kept relatively moist, root growth is reduced

which favors bulb enlargement and consequently flowering, since flower production in

bulbs is highly affected by bulb size in several species including Hippeastrum,

Habranthus, Zep,1iyiuhinhe (Theron and De Hertogh, 2001). A frequent and light

irrigation is commonly practiced on both vegetable and ornamental bulbous crops, while

over-irrigation is avoided since it can increase the incidence of several diseases (Hafeez,

1984).

Soil water deficits inhibit leaf expansion, as nutrient uptake is reduced because of

reduced transpiration rates (Russel, 1977). In onion, transpiration rates, photosynthesis

and growth are lowered by mild water stresses (Begun et al., 1990). Stressed onions often

bulb too early, produce small-sized bulbs and have an increased rate of bulb split. All

these factors can reduce marketable yields (Hegde, 1986).

The soil moisture requirement of onions is influenced by several factors including

cultivar used, soil type and temperature, light levels and other environmental factors. A

crop can be grown to maturity under a soil moisture deficit, but higher yields were









associated with irrigation frequencies that eliminated soil moisture deficits (De Lis et al.,

1967).

Minimizing the period when leaf and flower tissues are wet by using proper

irrigation management reduce moisture ad consequently the development of most fungal

and bacterial caused diseases. Fertilizer formulations also influence disease development

in flower bulb crops since they can alter soil pH, therefore fertigation should be carefully

planned in bulb production.

Nutrient requirements for greenhouse and outdoor bulb production can be grouped

into four categories according to De Hertogh and Le Nard (1993): 1) bulbs containing

sufficient nutrients to produce high quality potted plants or cut flowers without additional

fertilization such as Hippeastrum, Hyacinthus and Narcissus; 2) bulbs that require either

no additional fertilization or in which the application of Ca(N03)2 can eliminate or

reduce physiological disorders such as Dutch irises and tulips; 3) bulbs that require low

fertilization programs, such as Anemone, Freesia and Liatris; and 4) bulbs that require

moderate fertilization programs, such as Dahlia, Gladiolus, Lilium and Ranunculus.

Fertilization programs have been evaluated on several genera of flowering bulbs to

determine their effect on flowering, flower quality, bulb growth, bulb yield, and seed

yield. Flowering as well as bulb growth of Iris, Tulipa and Lilium were compared when

plants were treated with 10.7:3.9, 12.1:5.1, 14.3:3.9, or 17.9:3.9 meq/L of N and K. (Lee

et al., 2005). Results showed that flowering was slightly accelerated when Iris was grown

with 10.7:3.9 meq/L N:K; flowering of Tulipa was promoted by 14.3:3.9 meq/L N:K; and

neither growth or flowering of Lilium was significantly different between any fertilizer

treatments. When 0, 30, 70, 120, 180, 250, 330, 420, and 520 kg N ha-1 were applied to









Lachenalia cultivars, results demonstrated that increased rates of nitrogen had a positive

influence on bulb fresh mass and circumference (Engelbrecht, 2004). Three levels of

fertilizer, 50, 100, and 200 mg 1-1 N, were applied to Curcuma alismatifolia of the

Zingiberaceae family and results showed that plants treated with the highest fertilizer

level were taller, produced more stems per cluster, had larger diameter rhizomes, a

greater number of new rhizomes and better flower quality (Ruamrungsri et al., 2005).

In onions, irrigation intervals of once a week, every 10 days and every other week

were investigated in association with fertilizer levels of 90 and 180 kg N/ha to ascertain

the optimum irrigation interval and fertilizer level that produced the highest yields of

good quality bulbs. Results demonstrated that the 10 day irrigation interval and the 90 kg

N/ha rate resulted in the highest yield per ha and the greatest average bulb weight.

However, no treatment interactions were detected (Hassan, 1984). Irrigation frequencies

were also investigated on Caladium tubers in a pot study and results showed that three

times a week provided the best yield response compared to once a week and twice a week

(Overman and Harbaugh, 1988).

Extremely limited information was found in the horticultural literature regarding

Habranthus and Ze7,10/ii- uhe' fertigation frequencies or fertilizer regimes. The present

study was designed to address this issue and two experiments were conducted: 1) to

determine optimal fertigation frequency for these two genera in order to achieve an

extended flowering season with increased number of flowers per bulb, and 2) to

determine optimal fertilizer rates for bulb development.

In the first experiment, different fertigation frequencies (twice a week, once a week

and every other week) were applied to Habranthus robustus and Zephi7,/I/,wihe spp. to









determine their effect on flowering and/or on bulb development (number of leaves,

number of offsets, bulb size, bulb weight and number of buds). In the second experiment,

different fertilizer rates 0, 75, 150 and 300 ppm N, were applied to Habranthus robustus

and Ze7,1'/iiiwhe spp to verify their effect on flowering and bulb development.

The Habranthus species used in this project Habranthus robustus, is the most

popular species in the genus; it is native to Brazil but grows very well in Florida. The

Z.7,1'1/ riaihe species used in this projects were: Z. 'Paul Niemi', Z. JoAnn's Trial', and

Z. 'Fadjar Pink' which were hybridized by Fadjar Marta in Jakarta, Indonesia.

Experiment 1: Influence of three different fertigation treatments on number of

flowers and bulb development in Habranthus robstus and Ze7plh)1 ,iIhi spp.

Materials and Methods: Bulbs used in this experiment were obtained from

University of Florida stock, which had been grown in ground beds for a year prior to the

beginning of the experiment. On July 02, 2004, these bulbs were planted in 15 cm plastic

pots, three bulbs per pot using sphagnum peat based Fafard No. 2 soiless growing

medium (Agawam, MA) consisting of 70% Canadian sphagnum peat, 10% perlite and

20% vermiculite. All plants were placed in a greenhouse (Figure 4-1) under 11% shade,

natural photoperiods and a temperature range of 31/24C (day/night). These plants

received 250 ml of water with N at 150 mg.L-1, using Peters Professional 'Florida

Special', a water soluble fertilizer with 20N-4.7P-16.6K (Scotts Co., Marysville, OH).

This experiment was conducted in two different years: year 1 July 15 to

December 1, 2004 (week 29 to 47), and year 2 May 3 to December 1, 2005 (week 18 to

49). Fertigation treatments of 1) twice a week, 2) once a week, and 3) every other week

were started on July 15, 2004 the first year and May 3, 2005 on the second year.









There were 16 pots (48 bulbs) per treatment. During the growing phase of the

experiment, number of flowers was recorded twice a week during both years. A flower

was counted when the petals opened to expose the anthers and was removed after

collection of data. At completion of each year all bulbs were sectioned in order to

compare bulb development and floral initiation. Number of leaves, number of offsets

produced, total fresh weight (bulb with all leaves, roots and offsets attached), fresh

weight (bulb only), bulb size (diameter), and number of flower buds produced were

recorded, and were subjected to regression analysis.


















Figure 4-1: Habranthus bulbs during fertigation experiment with different plastic tags in
different color distinguishing the three treatments


























(A) (B)
Figure 4-2: Habranthus bulbs being weighed after completion of experiment, only bulb
(A) and bulb with leaves, roots and offsets (B)

Results and Discussion

The effects of the different fertigation treatments on number of flowers of

Habranthus robustus could not be analyzed during the first year as the data collection

started after the flowering peak for this crop.

During year 2 plants responded differently to treatments regarding the number of

flowers (Figure 4-3). Plants that were fertigated every other week flowered one week

earlier than plants fertigated once and twice a week. This may have occurred due to

drought stress. These plants ended their flowering season two weeks earlier than plants

fertigated once a week and eight weeks earlier than plants fertigated twice a week. Plants

fertigated once and twice a week started flowering during week 19 but plants fertigated

twice a week flowered longer, until week 37, while plants fertigated once a week stopped

flowering in week 31. Peak flowering occurred much earlier in plants fertigated twice a

week (week 21), than plants fertigated once a week (week 24) and every other week

(week 26). Number of flowers produced during peak flowering differed dramatically









among the treatments, as plants fertigated twice a week had 41 flowers compared to 25

for plants fertigated once a week and 34 for plants fertigated every other week.

Plants fertigated once and twice a week produced flowers more consistently than

plants fertigated every other week, as these plants had a flush of flowers at the beginning

of the season, flowered poorly during following weeks, did not flower for a month and

then had a second flush of flowers. Both regimens of fertigation watered twice or once

a week seemed to be effective on Habranthus under greenhouse environment, as these

regimens resulted in longer flowering periods and greatest number of flowers.

Similar results were observed in onions where water deficits resulted in

underdeveloped plants with reduced yields (Kadayifci et al., 2004). According to

Mermoud et al. (2005) changes in the irrigation frequency significantly influences the

components of the water balance. A decrease in irrigation frequency causes an increase in

the water storage in the root zone and a moisture deficit in the immediate vicinity of the

soil surface, which affect the crop's performance. This may have occurred in this study

when plants were fertigated every other week.

The results obtained after completion of the first year demonstrated that different

fertigation frequencies affected Habranthus bulb development. Regression analyses of all

factors investigated (number of leaves, number of offsets, bulb size, total fresh bulb

weight, bulb weight and number of flower buds) showed linear responses with coefficient

of determination above 0.50. All factors increased as fertigation increased in year 1

(Figures 4-4 to 4-9). Number of offsets, total fresh bulb weight and bulb weight had the

highest coefficient of determinations; all were above 0.90. These results can be compared

to those obtained by Overman and Harbaugh (1988) who tested different irrigation









frequencies in Caladium tubers and observed that most frequent irrigations resulted in the

best yields.

After completion of the second year the results obtained supported the conclusion

that different fertigation systems affect some aspects of Habranthus bulb development as

described bellow. There were no linear responses for number of leaves and number of

flower buds (Figures 4-10 and 4-15). Number of offsets decreased as fertigation

frequencies increased (r=l) (Figure 4-11). Bulb size, total fresh bulb weight and bulb

weight increased as fertigation frequencies increased; however bulb weight (total and

only bulb) had higher coefficients of determination; both were above 0.90 (Figures 4-12,

4-13 and 4-14).

During year 1 Zei7,y,1lit wuihe responded differently regarding flowering to different

fertigation frequencies compared to Habranthus. This was evident in Ze7,1h ,ainhe,\

during the first year, but not in Habranthus as both species (Habranthus and

Z7,' 1l/, lahe' ) have different flowering seasons and Z7,10yl-, iahei plants begin to flower

much later than Habranthus plants (Figure 4-16).

The results of year 1 showed that Ze7,1yl)-inwhe bulbs fertigated once a week and

twice a week had three gradual peaks of flower followed by an abrupt decline. This did

not happen when plants were fertigated every other week. Plants fertigated once and

twice a week flowered similarly which differed from plants fertigated every other week

(Figure 4-17).

Plants fertigated twice a week and every other week flowered for one week more

than the ones fertigated once a week. Plants fertigated twice a week had, overall, a greater









quantity of flowers during the entire season, but plants fertigated every other week had

more flowers during the first peak -37 flowers in a group of 36 bulbs (Figure 4-17).

During year 2 plants fertigated once a week and twice a week demonstrated three

flowering peaks followed by an abrupt decline in flower production. These two groups

had similar flowering patterns, which differed from plants fertigated every other week.

Plants fertigated twice a week had a greater number of flowers (Figure 4-18).

Plants fertigated once a week started flowering two weeks earlier than plants

fertigated twice a week and finished their flowering season one week before both plants

fertigated twice a week and every other week. Plants fertigated twice a week had, overall,

a greater number of flowers during the entire season, but plants fertigated every other

week had more flowers during their first peak -39 flowers in a group of 36 bulbs.

Plants that were fertigated once and twice a week produced their first flowers

earlier in the season (week 19 for plants fertigated once a week and week 21 for

fertigated twice a week), but plants under both regimens flowered poorly for more than

two months, until the first flush of flowers, on week 31.

Both once and twice a week fertigation regimens seemed to be effective for

flowering in Ze7,'/I )-ihe,' under greenhouse environments. But fertigating twice a week

would be preferable since the results showed that plants under this fertigation regimen

produced more flowers. Similar experiments were performed with onion and garlic (a

tunicate type and a non-tunicate type of bulb), and a once to twice a week watering

regimen has also been found to be preferable for these crops (Hanson et al., 2003;

Kadayifci et al., 2004; Mermoud et al. 2005). Fertigating plants every other week reduced

flowering of Zepi, ilie/,'\ bulbs as it resulted in a shorter flowering season with intervals









of no flowers. Similar experiments could be done in further studies using different

conditions of light and temperature in order to better define the ideal fertigation

frequency for these two species of tropical bulbs.

The treatments affected Z7p,10y Il n1hel bulb development in year 1. Regression

analysis of all factors investigated, with the exception of number of offsets (Figure 4-20),

showed linear responses with coefficient of determination above 0.50. All factors that had

a linear response (number of leaves, bulb size, total fresh bulb weigh, bulb weight and

number of flower buds) increased as fertigation increased during the first year (Figures 4-

19, 4-21, 4-22, 4-23, and 4-24), similarly to the Habranthus study. Number of leaves and

fresh bulb weight had highest coefficient of determination, both were above 0.90.

During year 2, number of offsets and total fresh bulb weight were the only factors

that had linear responses, with coefficient of determination above 0.50 (Figures 4-25 to 4-

30). Number of offsets decreased as fertigation frequencies increased (Figure 4-26),

while total fresh bulb weight increased as fertigation frequencies increased and was

highest when bulbs were fertigated once a week (Figure 4-28).












U Fertigated twice a week U Fertigated once a week 0 Fertigated every other week


18 19 20 21 22 23 24 25 26


27 28
Time


29 30 31 32 33 34 35 36 37


Figure 4-3: Number of flowers ofHabranthus robustus bulbs in year 2 as affected by
fertigation frequency (n=48). Bars represent standard errors.


1=every other week
2=once a week
3=twice a week


Mean
- Linear (Mean)

y = 0.561x + 4.3687
R2= 0.5711


Fertigation frequency


Figure 4-4: Number of leaves of Habranthus robustus bulbs in year 1 as affected by
fertigation frequency.


08





E
o06


z
04


02-


T


8
S6
S4
S2
E


A -


T-


I













S=every other week
2=once a week
3=twice a week


4
4


0 1 2 3

Fertigation frequency


Mean

--Linear (Mean)

y =2.9065x- 1.204
R2= 0.9377


Figure 4-5: Number of offsets ofHabranthus robustus bulbs in year 1 as affected by
fertigation frequency.


345
34
335
33
325
32
3 15
0 1 2 3 4


= every other week
2=once a week
3=twice a week


Mean
- Linear (Mean)


y = 0.0845x + 3.1387
R2= 0.8541


Fertigation frequency


Figure 4-6: Bulb size ofHabranthus robustus in year 1 as affected by fertigation
frequency.


* Mean


4


1=every other week
2=once a week
3=twice a week


Linear

y = 6.7565x + 32.339
R2= 0.9919


Fertigation frequency


Figure 4-7: Total weight ofHabranthus robustus bulbs in year 1 as affected by fertigation
frequency.


ci,
8n
o 6
04
2,
E o
D
Z


60
40
(,
20
5 o


0








76



1 =every other week
2=once a week
3=twice a week
25
Mean
"2 -- Linear (Mean)
d 15
10
Sy= 1.8475x+ 14.411
R2 = 0.9958
0
0 05 1 15 2 25 3 35
Fertigation frequency


Figure 4-8: Bulb weight ofHabranthus robustus in year 1 as affected by fertigation
frequency.


1=every other week
2=once a week
3=twice a week
25
0 2

5 15 Mean
S1 Linear (Mean)
Z 05-
0z ___ y =0.236x + 1.3603
0 05 1 15 2 25 3 35 R2 = 0.604
Fertigation frequency


Figure 4-9: Number of flower buds of Habranthus robustus bulbs in year 1 as affected by
fertigation frequency.


= every other week
2=once a week
7_ 3=twice a week
6
5 5 Mean
4- Linear (Mean)
3-
2- y = 0.2395x + 4.936
Z R2 = 0.1401
0-
0 05 1 15 2 25 3 35
Fertigation frequency


Figure 4-10: Number of leaves ofHabranthus robustus bulbs in year 2 as affected by
fertigation frequency.













S=every other week
2=once a week
3=twice a week


* Mean
- Linear (Mean)

y = -0.375x + 19.75
R2=1


0 05 1 15 2 25 3 35
Fertigation frequency


Figure 4-11: Number of offsets ofHabranthus robustus bulbs in year 2 as affected by
fertigation frequency.


= every other week
2=once a week
3=twice a week


Mean
- Linear (Mean)

y =0.1155x +2.8087
R2 = 0.5937


0 05 1 15 2 25 3 35
Fertigation frequency


Figure 4-12: Bulb size ofHabranthus robustus in year 2 as affected by fertigation
frequency.


70
60
50
") 40
S30
S20
10
0


0 05 1 15 2
Fertigation frequency


1=every other week
2=once a week
3=twice a week


Mean
- Linear (Mean)


y = 6.777x + 37.928
R2 = 0.9685


25 3 35


Figure 4-13: Total bulb weight ofHabranthus robustus in year 2 as affected by
fertigation frequency.


















15

10

5-

0
0 05 1 15 2 25 3 35
Fertigation frequency


S=every other week
2=once a week
3=twice a week


* Mean
Linear (Mean)


y =3.0715x + 8.695
R2 = 0.9973


Figure 4-14: Bulb weight of Habranthus robustus in year 2 as affected by fertigation
frequency.


S=every other week
2=once a week
3=twice a week


Mean
- Linear (Mean)

y = 0.0855x + 2.7053
R2 = 0.3893


0 05 1 15 2 25 3 35
Fertigation frequency


Figure 4-15: Number of flower buds ofHabranthus robustus bulbs in year 2 as affected
by fertigation frequency.


305
3
n 295
o 29
285
E 28
Z 275
27












-*-Total flw Habrabthus Total flwZephyranthes


Time
0
60
E
z

40




20



0
26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47
Time

Figure 4-16: Total number of flowers on Habranthus and 7Zep2lyt unlthe bulbs from July
to December 2004 (n=142)


U Fertigated tAwce a week U Fertigated once a week O Fertigated every other week


14-


12






108-
1


,-r


z


02-


0


31 32 33 34 35 36 37 38 39 40 41 42 43 44
Week


45 46


Figure 4-17: Number of flowers of Zepian/)theS' spp. bulbs in year 1 as affected by
fertigation frequency (n=48). Bars represent standard errors.


STTT T





T T l l Ii ,








80






U Fertigated twice a week U Fertigated once a week 0 Fertigated every other week


19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46
Week

Figure 4-18: Number of flowers of Ze7,I,'/l i ies spp. bulbs in year 2 as affected by
fertigation frequency (n=48). Bars represent standard errors.


4
3
2
1
0-


0 05 1 15 2
Fertigation frequency


25 3 3


S=every other week
2=once a week
3=twice a week


Mean
- Linear (Mean)


y =1.0155x + 1.6587
5 R2 = 0.9942


Figure 4-19: Number of leaves of Zepjy i,,1 i/he spp. bulbs in year 1 as affected by
fertigation frequency.


12

. 1
u=

U 08
E
0
z
06

04

02

n


T-

T -- t l -

















8
97
6
95
4
3
2

9
0 05 1 15 2 25 3 35
Fertigation frequency


Figure 4-20: Number of offsets of Zly 1,j)i ilh,\
fertigation frequency.


0 05 1 15 2
Fertigation frequency


S=every other week
2=once a week
3=twice a week


Mean
- Linear (Mean) 3

R2= 0.1242


spp. bulbs in year 1 as affected by


1=every other week
2=once a week
3=twice a week


Mean
--Linear (Mean)


y =0.0725x + 2.2127
R2 = 0.7707


25 3 35


Figure 4-21: Bulb size of Zephi) /,1il/l spp. in year 1 as affected by fertigation
frequency.


339
338
33 7
336
335
334
33 3
332
331
33 -
329


0 05 1 15 2
Fertigation frequency


S=every other week
2=once a week
3=twice a week


Mean
Linear (Mean)

y = 0.3455x + 32.756
R2 = 0.7381


25 3 35


Figure 4-22: Total fresh weight of Zey7,ph/ill/h\ spp. bulbs in year 1 as affected by
fertigation frequency.















1315
131
1305
13
1295
129


0 05 1 15 2 25 3 35
Fertigation frequency


Figure 4-23: Bulb weight of Zeply',t)/iih, spp. in year 1
frequency.


S=every other week
2=once a week
3=twice a week
Mean
--Linear (Mean)


y = 0.08x + 12.877
R2 = 0.9796




as affected by fertigation


1=every other week
2=once a week
3=twice a week


0 05 1 15 2
Fertigation frequency


* Mean
--Linear (Mean)


y =0.196x + 2.0313
R2= 0.6214


25 3 35


Figure 4-24: Number of flower buds of Zep li'/i/,1trh spp. bulbs in year 1 as affected by
fertigation frequency.


1=every other week
2=once a week
3=twice a week


Mean
--Linear (Mean)

y = -0.0275x + 3.8513
R2 = 0.0237


Fertigation frequency


Figure 4-25: Number of leaves of Z7 ,1 w ,i /rithe
fertigation frequency.


spp. bulbs in year 2 as affected by


0)
a), 4-
S395
- 39
0 3 85
, 38-
E 375
3 37-
z R_









1=every other week
2=once a week
3=twice a week


Mean
--Linear (Mean)

y =-0.5835x +10.111
R2 = 0.6446


0 1 2 3 4
Fertigation frequency

Figure 4-26: Number of offsets of Ze7,/phl iu(/l spp. bulbs in year 2 as affected by
fertigation frequency.


*
0 1 2 3 4
Fertigation frequency


Figure 4-27: Bulb size of Zepy,1itillhe
frequency.


= every other week
2=once a week
3=twice a week
Mean
- Linear (Mean)

y = 0.0205x + 2.6507
R2 = 0.3058


spp. in year 2 as affected by fertigation


= every other week
2=once a week
3=twice a week


Mean
--Linear (Mean)
y = 0.85x + 34.4
R2 = 0.6628


Fertigation frequency

Figure 4-28: Total fresh weight of Ze7,1phyt)I/il spp. bulbs in year 2 as affected by
fertigation frequency.


i


i"







84






13 05 0 ________ ____________________ *ean
13 05

12 95 Mean
0 12 9
S12 85 Linear (Mean)
;Q 12 85 ------ -k i-------------
S 12 8
m 1275
12 7
1265 y = 0.09x + 12.657
0 1 2 3 4
R2 = 0.3166
Fertigation frequency


Figure 4-29: Bulb weight of Zeil,) i//,whe% spp. in year 2 as affected by fertigation
frequency.





2 575
S257 Mean
2565 Linear (Mean)
256
2 555
:3 2555 ---- -^"-----------
S2 55 y = 0.005x + 2.55
2545 R2 = 0.25
0 1 2 3 4
Fertigation frequency


Figure 4-30: Number of flower buds of Zey),1 i t, u/lie spp. bulbs in year 2 as affected by
fertigation frequency.

Experiment 2: Effect of different fertilizer rates on leaf production and bulb


development in Habranthus robustus and Ze7,1y t awihe\ spp.


Materials and Methods: On November 6, 2005, 36 Habranthus robustus and


Ze7,1'l)o -1 he, spp. bulbs were selected from University of Florida stock, which had been


grown in ground beds for two years prior to the beginning of this study. Each bulb had


existing bulblets (offsets) removed, two leaves per Ze7,1y ,l) n1he, bulb and four leaves per


Habranthus (extra leaves were also removed). All bulbs were weighed and the diameter


measured prior to planting (Table 4-1).