Citation
Root-zone temperature and soil moisture effects on growth and physiology of container-grown 'Grande Naine' banana and Ixora chinensis L. 'Maui'

Material Information

Title:
Root-zone temperature and soil moisture effects on growth and physiology of container-grown 'Grande Naine' banana and Ixora chinensis L. 'Maui'
Creator:
Ramcharan, Christopher, 1943-
Publication Date:
Language:
English
Physical Description:
xii, 172 leaves : ill. ; 28 cm.

Subjects

Subjects / Keywords:
Bananas ( lcsh )
Dissertations, Academic -- Horticultural Science -- UF
Horticultural Science thesis Ph. D
Ixora chinensis ( lcsh )
Plant roots ( jstor )
Photosynthesis ( jstor )
Human growth ( jstor )
Genre:
bibliography ( marcgt )
non-fiction ( marcgt )

Notes

Thesis:
Thesis (Ph. D.)--University of Florida, 1987.
Bibliography:
Bibliography: leaves 160-171.
General Note:
Typescript.
General Note:
Vita.
Statement of Responsibility:
by Christopher Ramcharan.

Record Information

Source Institution:
University of Florida
Holding Location:
University of Florida
Rights Management:
Copyright [name of dissertation author]. Permission granted to the University of Florida to digitize, archive and distribute this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
Resource Identifier:
030394745 ( ALEPH )
16959530 ( OCLC )
AER3637 ( NOTIS )
AA00004845_00001 ( sobekcm )

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Full Text


ROOT-ZONE TEMPERATURE AND SOIL MOISTURE EFFECTS
ON GROWTH AND PHYSIOLOGY
OF CONTAINER-GROWN 'GRANDE NAINE' BANANA
AND IXORA CHINENSIS L. 'MAUI'
By
CHRISTOPHER RAMCHARAN
A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA
IN PARTIAL FULFILLMENT OF THE REQUIREMENTS
FOR THE DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1987


ACKNOWLEDGEMENTS
I want to especially thank my major advisor, Dr.
Dewayne. L. Ingram. He impressed upon me the art of
scientific writing, an enthusiasm for research and the will
to persevere. Also, I wish to thank the other members of my
committee, Drs. Terril Nell, James Barrett, Jerry Bennett and
William Wiltbank, for their encouragement and critical
evaluation of my work.
I am deeply grateful to the United States Department of
Agriculture whose financial support and grant made this
research possible and to the Department of Ornamental
Horticulture for support staff, equipment and services. The
contributions of Ms. Claudia Larsen for laboratory and
technical assistance and T.D. Townsend of the greenhouse
staff are highly appreciated.
I would like to thank Oglesby nursery for the generous
donations of banana plants and Mr. George Behren of Behren's
nursery for the ixora plants used in this study.
Last, but certainly not least, sincere thanks go to my
family. The support of my wife Senovia throughout the course
of these studies was invaluable. Finally, I wish to
acknowledge my children: they gave me the purpose for which
it was done.


TABLE OF CONTENTS
Page
ACKNOWLEDGEMENTS ii
LIST OF TABLES v
LIST OF FIGURES vii
ABSTRACT xi
CHAPTERS
I GENERAL INTRODUCTION 1
II LITERATURE REVIEW 4
Introduction 4
Water Stress Physiology 5
Physiological Effects of Root-Zone
Temperature 12
Root-Zone Temperature and Soil Moisture
Interactions 18
IIIEFFECTS OF IRRIGATION VOLUME ON BANANA
AND IXORA UNDER TWO GROWING CONDITIONS 22
Introduction 22
Materials and Methods 23
Plant Materials and General Cultural
Procedures 23
Greenhouse 23
Experiment 1 23
Experiment 2 26
Growth Room 26
Experiment 3 26
Results and Discussion 27
Greenhouse 27
Experiment l--banana 27
Experiment l--ixora 34
Experiment 2--banana 40
Experiment 2--ixora 42
Growth Room 42
Experiment 3--banana 42
Experiment 3--ixora 50
i i i


IVROOT-ZONE TREMPERATURE EFFECTS ON BANANA AND
IXORA UNDER TWO GROWING CONDITIONS 56
Introduction 56
Materials and Methods 57
Plant Materials and General Cultural
Procedures 57
Experiment l--Greenhouse 57
Experiment 2--Growth Room 60
Results and Discussion 61
Experiment l--Greenhouse 61
Banana 61
Ixora 65
Experiment 2--Growth Room 68
Banana 68
Ixora 72
VROOT-ZONE TEMPERATURE AND SOIL MOISTURE
EFFECTS ON BANANA AND IXORA UNDER TWO
GROWING CONDITIONS 77
Introduction 77
Materials and Methods 78
Plant Materials and General Cultural
Procedures 78
Experiment l--Greenhouse 79
Experiment 2--Growth Room 84
Results and Discussion 86
Experiment l--Greenhouse 86
Physiological responses--banana .... 86
Growth responses--banana 93
Carbohydrate analysis--banana 100
Physiological responses--ixora 104
Growth responses--ixora 109
Carbohydrate analysis--ixora 118
Experiment 2--Growth Room 120
Physiological responses--banana .... 120
Growth responses--banana 128
Carbohydrate ana 1ysis--banana 132
Physiological responses--ixora 137
Growth responses--!* xora 145
Carbohydrate analysis--ixora 150
VISUMMARY AND IMPLICATIONS 153
LITERATURE CITED 160
BIOGRAPHICAL SKETCH 172


LIST OF TABLES
Page
5-1. Growth components of 'Grande Naine' banana
measured after 10 weeks at four root-zone
temperatures and two irrigation volumes under
greenhouse conditions 94
5-2. Dry weight components of 'Grande Naine' banana
measured after 10 weeks at four root-zone
temperatures and two irrigation volumes under
greenhouse conditions 98
5-3. Shoot and root carbohydrate distribution of
'Grande Naine' banana at four root-zone
temperatures and two irrigation volumes under
greenhouse conditions 101
5-4. Growth components of Ixora chinensis L.'Maui*
measured after 10 weeks at four root-zone
temperatures and two irrigation volumes under
greenhouse conditions 112
5-5. Shoot and root carbohydrate distribution of
Ixora chinensis L. 'Maui' measured after 10 weeks
at four root-zone and two irrigation volumes
under greenhouse conditions 119
5-6. Growth components of 'Grande Naine' banana
measured after six weeks at four root-zone
temperatures and two irrigation volumes under
growth room conditions 129
5-7. Dry weight components of 'Grande Naine' banana
measured after six weeks at four root-zone
temperatures and two irrigation volumes under
growth room conditions 130
5-8. Shoot and root carbohydrate distribution of
'Grande Naine' banana at four root-zone
temperatures and two irrigation volumes under
growth room conditions 134
v


5-9. Growth components of Ixora chinensis L.'Maui*
measured after six weeks at tour root-zone and
two irrigation volumes under growth room
conditions 146
5-10. Dry weight components of Ixora chinensis L.'Maui'
measured after six weeks at four root-zone
temperatures and two irrigation volumes under
growth room conditions 147
5-11. Shoot and root carbohydrate distribution of
Ixora chinensis L. 'Maui' measured after six weeks
at four root-zone temperatures and two irrigation
volumes under growth room conditions 151


LIST OF FIGURES
Page
3-1. Photosynthetic photon flux density and relative
humidity in the greenhouse during measurements
of physiological parameters 24
3-2. Effects of three irrigation volumes on the
diurnal physiological responses of 'Grande
Naine' banana under greenhouse conditions 29
3-3. Effects of three irrigation volumes on the
diurnal water use efficiency of 'Grande Naine'
banana under greenhouse conditions 33
3-4. Effects of three irrigation volumes on the
diurnal physiological responses of Ixora
chinensis L. 'Maui' under greenhouse
conditions 36
3-5. Effects of three irrigation volumes on the
diurnal water use efficiency of Ixora
chinensis L. 'Maui' under greenhouse
conditions 39
3-6. Effects of a 14 day drying cycle on leaf water
potential and leaf conductance of 'Grande Naine'
banana under greenhouse conditions 41
3-7. Effects of a 14 day drying cycle on leaf water
potential and leaf conductance of Ixora
chinensis L. 'Maui'under greenhouse
conditions 43
3-8. Effects of three irrigation volumes on the
diurnal physiological responses of 'Grande Naine'
banana under growth room conditions 45
3-9. Effects of three irrigation volumes on the
diurnal water use efficiency of 'Grande Naine'
banana under growth room conditions 49
vi 1


3-10. Effects of three irrigation volumes on the
diurnal physiological responses of Ixora
chinensis L. 'Maui' under greenhouse
conditions 52
3-11. Effects of three irrigation volumes on the
diurnal water use efficiency of Ixora
chinensis L. 'Maui under growth room
conditions 55
4-1. Photosynthetic photon flux density and relative
humidity in the greenhouse during measurements of
physiological parameters 58
4-2. Effects of four root-zone temperatures on the
diurnal physiological responses of 'Grande
Naine banana grown under greenhouse
conditions 63
4-3. Effects of four root-zone temperatures on the
diurnal physiological responses of Ixora
chinensis L. 'Maui' grown under greenhouse
conditions 67
4-4. Effects of four root-zone temperatures on the
diurnal physiological responses of 'Grande
Naine' banana grown under growth room
conditions 70
4-5. Effects of four root-zone temperatures on the
diurnal physiological responses of Ixora
chinensis L. 'Maui' grown under growth room
conditions 74
5-1. Photosynthetic photon flux density and relative
humidity in the greenhouse during measurements of
physiological parameters 80
5-2. Effects of four root-zone temperatures and a 50+_5
ml daily irrigation volume per 555 cm3 container
on the diurnal physiological responses of 'Grande
Naine' banana grown under greenhouse
conditions 88
5-3. Effects of four root-zone temperatures and a 100+_10
ml daily irrigation volume per 555 cm3 container
on the diurnal physiological responses of 'Grande
Naine' banana grown under greenhouse
conditions 90
5-4. Regressions of midday physiological responses of
container-grown 'Grande Naine' banana over
four root-zone temperatures and two irrigation
volumes under greenhouse conditions 92
v i i i


5-5. Effects of four root-zone temperatures across
two irrigation volumes on leaf area of the third
newest leaf in container-grown 'Grande Naine'
banana measured over 10 weeks under greenhouse
conditions 96
5-6. Effects of four root-zone temperatures and two
irrigation volumes on shoot/root ratio and root
dry weight of container-grown 'Grande Naine'
banana grown under greenhouse conditions 99
5-7. Effects of four root-zone temperatures and two
irrigation volumes on root sugar/starch content
of container-grown 'Grande Naine' banana
under greenhouse conditions 103
5-8. Effects of four root-zone temperatures and a 50+_5
ml daily irrigation volume per 555 cm3 container
on the diurnal physiological responses of Ixora
chinensis L. 'Maui' grown under greenhouse
conditions 106
5-9. Effects of four root-zone temperatures and a 100+_10
ml daily irrigation volume per 555 cm3 container
on the diurnal physiological responses of Ixora
chinensis L. 'Maui' grown under greenhouse
conditions 108
5-10. Regressions of midday physiological responses of
container-grown Ixora chinensis L. 'Maui' over
four root-zone temperature and two irrigation
volumes under greenhouse conditions Ill
5-11. Main effects of four root-zone temperatures
across two irrigation volumes on growth of Ixora
chinensis L. 'Maui' grown under greenhouse
conditions 115
5-12. Effects of four root-zone temperatures and two
irrigation volumes on dry weight components of
container-grown Ixora chinensis L. 'Maui'
grown under greenhouse conditions 117
5-13. Effects of four root-zone temperatures and a 75+_8
ml daily irrigation volume per 1200 cm3 container
on the diurnal physiological responses of
container-grown 'Grande Naine' banana grown under
growth room conditions 122
5-14. Effects of four root-zone temperatures and a 150 +_15
ml daily irrigation volume per 1200 cm3 container
on the diurnal physiological responses of container-
grown 'Grande Naine' banana grown under
growth room conditions 124
i x


5-15. Regressions of midday physiological responses
of container-grown 'Grande Naine' banana over
four root-zone temperatures and two irrigation
volumes under greenhouse conditions 127
5-16. Effects of four root-zone temperatures across
two irrigation volumes on chlorophyll
concentration of container-grown 'Grande Naine'
banana under growth room conditions 133
5-17. Interactive effects of four root-zone temperatures
and two irrigation volumes on carbohydrate content
of container-grown 'Grande Naine' banana
under growth room conditions 137
5-18. Effects of four root-zone temperatures and a 75+8
ml daily irrigation volume per 1200 cm3 container
on the diurnal physiological responses of
Ixora chinensis L. 'Maui' grown under growth
room conditions 140
5-19. Effects of four root-zone temperatures and a 150+15
ml daily irrigation volume per 1200 cm3 container
on the diurnal physiological responses of
Ixora chinensis L. 'Maui' grown under growth
room conditions 142
5-20. Regressions of midday physiological responses
of container-grown Ixora chinensis L. 'Maui'
over four root-zone temperatures and two
irrigation volumes under growth room
conditions 144
5-21. Effects of four root-zone temperatures and two
irrigation volumes on growth of container-grown
Ixora chinensis L. 'Maui' under growth room
condi tions 149
x


Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy
ROOT-ZONE TEMPERATURE AND SOIL MOISTURE EFFECTS ON GROWTH
AND PHYSIOLOGY OF CONTAINER-GROWN 'GRANDE NAINE'
BANANA AND IXORA CHINENSIS L. 'MAUI'
By
Christopher Ramcharan
May 1987
Chairman: Dewayne L. Ingram
Major Department: Horticultural Science
Studies to investigate the effects of root-zone
temperature (RZT) and irrigation volume (IRV) on growth and
physiological responses of 'Grande Naine' banana (Musa spp.
AAA) plantlets and Ixora chinensis L. 'Maui' were conducted
under greenhouse (GH) and growth room (GR) environments.
Leaf photosynthesis (PS), conductance (CS) and
transpiration (TR) in the GH were reduced 65%, 57% and 49%,
respectively, in banana under an IRV of 20+_4 ml daily per 150
cm3 container as compared with the highest IRV of 40+8 ml
daily per container. Measured parameters were severely
reduced at an IRV of 10+2 ml per container. Physiological
responses of ixora were also reduced by decreased IRV. Leaf
wilting occurred at a leaf water potential (LWP) of -1.1 MPa
and leaves abscised below a LWP of -2.1 MPa.
xi


In an RZT study, maximum midday PS in banana occurred at
the 33C RZT in a GH and a maximum midday PS of 0.74 mg CO2
m"2 s'1 was observed in 33 and 38C RZT-treated plants in a
GR. In ixora, a 33C RZT induced a maximum midday PS in the
GH but there were no apparent differences in PS between 28,
33 and 38C RZTs in the GR study.
Maximum PS in banana occurred at 38 and 33C RZTs under
a higher IRV of 100+_10 ml daily in the GH. In the GR, maximum
PS occurred in 33C RZT-treated plants and the higher IRV but
at 38C RZT under a moderate IRV (75+8 ml daily). Increasing
RZT reduced leaf area under both IRVs. Carbohydrate (CHO)
partitioning between shoot and root was correlated with RZT-
induced changes in LWP.
Gas exchange processes in ixora were reduced by RZTs
above 33C RZT regardless of IRV. RZT-induced shoot growth
was not associated with increased PS or TR but a hormonal
role was theorized. CHO partitioning was likely related to
the induced shoot growth and was influenced by environmental
condi ti on s.
These findings could have significant applications to
banana production schemes using tissue-cultured plants. RZTs
above 33C could reduce the landscape and floricultural value
of container-grown ixora.
xi i


CHAPTER I
GENERAL INTRODUCTION
Temperature and water are perhaps two of the most
important environmental factors affecting growth and
physiology of plants. Although soil moisture can be modified
by irrigation, altering temperature is more difficult.
Extreme temperatures produce deleterious but often subtle
effects. Soil temperature and water are interrelated since
both directly affect plant roots. Maintaining optimum water
relations and temperature may be difficult, especially with
container-grown species.
Hot and dry conditions often occur simultaneously in
crop situations and alleviating the latter is often
considered a remedy for the former. Although many drought
tolerant plants are heat tolerant (85), the interactions
between temperature and water stresses have not been well
defined (102). Suboptimal root-zone temperatures (RZT) stress
plants through reduced respiration, increased water viscosity
(82) and reduced absorption caused by decreased cell membrane
permeability (74). In this context, increased RZTs, within
limits, are believed to have the reverse effect and so
increase water (5,30) and possibly nutrient absorption
(39,44).
1


2
Within limits, increased soil temperatures have
generally increased shoot growth curvi1inear 1y (30).
Restricted shoot or leaf elongation is usually the first
symptom of mild water stress (57). Reduced soil water might
offset increased shoot growth induced by increasing soil
temperature to within the optimum range. A similar induction
of increased nutrient absorption by increasing soil
temperature may occur thus permitting less than normal
nutrient application. Such a recommendation was made for
peppers (Capsicum annuum L.) grown at a supraoptimal RZT of
360c (44).
Water stress, by its immediate effect on shoot growth
but delayed action on photosynthesis (13), has been shown to
increase shoot carbohydrate concentrations (8,119). Increased
root temperature has also been shown to reduce shoot
carbohydrates (7), again suggesting the possibility of
interactive effects between the two stress factors. Indeed
some pasture grasses were of better quality because of higher
carbohydrate levels under high temperature and dry conditions
as opposed to being irrigated (18).
Interactive effects of soil water and RZT on
translocation and carbohydrate partitioning patterns have
been reported (7,8). Increased RZTs have altered
translocation rates due to increased sink demands of roots
(46). Water stress could reduce translocation by its effect
on reduced turgor pressure and the consequent effect on
decreased mass flow of assimilates.


Therefore, the overall objective of this study was to
identify the effects of supraoptimal RZT and water deficits
on two container-grown plants. The specific objectives were
to
1) Determine growth and physiological effects on 'Grande
Naine' banana (Musa spp. AAA) and ixora (Ixora chinensis L.
'Maui') and to elucidate whether the two stress factors act
independently or interactively.
2) Identify any compensatory type interactions that may
be exploited in container production of plants.
3) Determine if drought stress can be induced by high
RZT, identify critical RZTs and observe whether watering
offsets temperature-induced water stresses.
Banana and ixora were selected because they represent a
contrast in morphology, phenotype and horticultural usage.
The use of tissue-cultured banana plants in this study
represents a unique application of micropropagation as a tool
in plant physiology. The choice of banana was also in
anticipation of the stress problems that may arise in the
banana production schemes that convert to the use of tissue-
cultured plantlets.


CHAPTER II
LITERATURE REVIEW
Introduction
Temperature and water stress affect plant growth and
physiological processes. The literature is abundant with data
describing their independent effects but there have been few
attempts to separate the effects of these two stress factors
when applied in combination. Any attempt to determine
interactions between soil water and temperature on plants
must first review or determine their independent effects.
While it is generally accepted that water stress reduces
expansive growth and induces a progressive reduction in gas
exchange processes, the effects of supraoptimal RZTs have not
been clearly identified. Soil water deficits usually reduce
plant water status and supraoptimal RZTs may affect root
absorption and conductance and indirectly stomatal function.
However, plant water stress can occur even with adequate soil
water when ambient conditions favor excessive transpiration
or water absorption is inhibited by factors including high
RZT.
Since supraoptimal root temperature directly affects
root respiration and metabolism, its effect on partitioning
4


5
patterns within the plant seems likely more direct than that
of water stress.
Water stress physiology
Water stress basically results from an imbalance
between soil water absorption and that required by plants to
meet transpirationa1 needs. Kramer (74) stated that plant
water stress is caused by either excessive water loss,
inadequate absorption or a combination of the two. In
general, an increase in water stress produces a two phase
response in photosynthesis, with a threshold water potential
above which there is little or no change in photosynthesis
and below which the rate decreases rapidly (114). A water
potential of -0.4 MPa reduced the photosynthetic capacity of
loblolly pine with assimilation being negligible at -1.1 MPa
(17). This effect was caused by stomatal closure which
reduced transpiration as well as photosynthesis. Water stress
may affect photosynthesis via three major mechanisms (131):
1) by stomatal closure, 2) by cellular dehydration which
reduces enzyme activity, and 3) by decreasing cell membrane
permeability to carbon dioxide and bicarbonate.
Although stomatal closure has been cited as the primary
method by which water stress affects photosynthesis
(13,78,132), there are several reports of non-stomatal
reductions in photosynthesis (14,23,49,114). Many species
that exert some degree of stomatal limitation on
photosynthesis at low water potentials characteristically
have high mesophyll resistance (50,141). When the components


6
of leaf resistance to CO2 were measured for several species,
mesophyll resistance was found to be lower than stomatal
resistance but about ten times that of boundary layer
resistance (139). Soil water deficit also affects the diurnal
pattern of stomatal aperture (33) with progressi ve 1y earlier
closing under prolonged drought conditions (93).
Stomatal closure affected photosynthesis mainly by its
effect on the carbon dioxide supply (13,17). However, some
researchers agree that photosynthesis is evident even when
stomata are closed (93). Because of their diffusive nature,
photosynthesis and transpiration are both affected by
stomatal closure, although transpiration may be affected more
than photosynthesis (48,120). Photosynthesis was reduced 25%
while transpiration decreased 50% in apple (Mal us pumi1 a
Mill.) under drought conditions (51). Zelitch (149) indicated
that partial stomatal closure reduced transpiration
relatively more than photosynthesis. Although the diffusive
resistance of CO2 was larger than that of water, CO2 also
encountered a vastly greater mesophyll resistance than water.
Consequently, the stomata represented a smaller portion of
the total resistance to CO2 than to water vapor.
Moorby et al. (95) found no effect of drought on the
activity of ribulose 1,5-diphosphate (RUDP) carboxylase or
carbonate dehydratase, two enzymes involved in CO2 fixation.
Ackerson et al. (1) however, reported that water stress
reduced the activity of photosynthetic enzymes of potato
(Solanum tuberosum L). Lawlor (84) indicated that ATP


7
synthesis through photophosphorylation was inhibited by water
stress while electron transport and reductant supply were
relatively insensitive. Boyer (14) demonstrated that
photosynthesis of water stressed sunflower (He!ian thus annuus
L.) plants under growth chamber conditions was affected more
by photochemical activity than by CO2 diffusion or enzymatic
activity. Davies (33) showed that stomatal sensitivity to
decreasing leaf water potential in cotton (Gossypi urn hirsutum
L.) and soybean (Glycine max. L.) was greater for plants
grown in a growth chamber than in a greenhouse.
When soil moisture is adequate, diurnal fluctuations in
xylem pressure potential closely followed the daily trend of
atmospheric evaporative demand, whether expressed as
radiation, vapor pressure or humidity (22,71,121). As soil
moisture becomes limiting, early morning leaf water
potentials (LWP) begin at a lower value, decrease sharply and
then remain constant over several hours, reflecting the
influence of increased stomatal resistance (29,108).
Threshold relationships between diffusive resistance and leaf
water potential appear to be more common than linear or
curvilinear relationships (52,67,68). Brix (17) showed that
net assimilation rate (NAR) was closely related to LWP; NAR
began to decrease at -1.0 MPa and completely ceased at -3.5
MPa. Other reports (53,79,113) indicated that parallel
decreases in NAR and LWP were strongly linked to changes in
diffusive resistance. However, NAR changes in pear (Pyrus
communis L.) (77), Pacific silver fir (Abies alba Mill.) and


8
noble fir (Abies nobi 1 is Lindl.) (55) did not always coincide
with changes in diffusive resistance indicating the influence
of mesophyll and or carboxylation resistance. It was
concluded that species vary considerably in their
physiological reactions to water deficits.
The majority of studies on water stress in banana have
dealt primarily with the effects on fruit bunch and yield
characteristics, but with few details reported on
morphological, phenological and physiological responses.
However, all reports agree that water stress causes a general
reduction in growth (19,111,136). Daniells (31) in Australia
indicated a dramatic reduction in growth, delayed bunch
emergence, inferior fruits and longer maturity time for
'Williams' banana under decreasing soil water levels.
Robinson and Albertson (116) reported that optimum growth and
yields were obtained by maintaining soil moisture levels at
16 to 34% of total available soil moisture (ASM).
Bhattacharyya and Rao (10) also reported optimum bunch and
yield character!' sti cs with 20 to 40% ASM depletion levels. An
unimpeded supply of water was essential for development of an
efficient root system, rapid rate of leaf emergence,
continued production and partitioning of assimilates, and
synthesis of growth hormones such as cytokinins. In Honduras,
Ghavami (40) found an irrigation rate of 44 mm/week or a soil
moisture tension of 30 to 40 centibars resulted in the
highest yields of 'Valery' banana.


9
Perhaps the most detailed description of the effects of
soil water levels on banana physiology was recorded by
Shmueli (126) in his paper on the physiological activity of
banana in relation to soil moisture in Israel. Depletion of
soil moisture below a third of the available water capacity
was accompanied by marked yellowing of leaves and significant
reduction in stem diameter. Diurnal changes in stomatal
activity were clearly related to ASM. When soil moisture
decreased below two-thirds of total ASM, there was a
reduction in stomatal conductance which was confined mainly
to the morning hours. The impairment of plant water balance
became acute when soil moisture decreased to about one-third
of the total available. Although diurnal patterns of stomatal
conductance and transpiration generally paralleled each
other, the rate of transpiration was considerably more
influenced by environmental conditions. Soil moisture levels
below two-thirds of field capacity resulted in reduced leaf
osmotic potential. Overall, banana plants responded to
diminishing soil moisture by stomatal closure, reduced
assimilation and accelerated yellowing of leaves indicative
of starvation effects.
Current efforts to increase agricultural production,
coupled with conservation of limited high quality water, have
focused attention on the possibility of increasing water use
efficiency (WUE) of various crops (16,88,97). Fischer and
Turner (38) observed that under field conditions, WUE usually
remained constant or increased slightly with increasing soil


10
water limitations for C3 species. Reports on the effects of
water stress on WUE are conflicting. Some investigations
pointed to only small increases in WUE with water stress
(16,38,97). Reduction of water supply to wheat
(Triticum aestivum L.) such that transpiration was reduced by
70% of the controls was associated with a 20% increase in WUE
(41). On the other hand, WUE in maize (Zea mays L.) was lower
on days when there was increasing soil water stress (128).
Limited soil water resulted in yield losses but enhanced WUE
in grapefruit (Citrus paradisi L.) (16) and Shamouti orange
(Citrus sinensis Osb.) (97). Manning et al. (86) found a
positive correlation of WUE with soil moisture regime, plant
height, leaf area and seed yield in pea [Pisum sativum (L.)
Mer. ].
Adequate irrigation was critical for vegetative growth
in coffee (Coffea arabica L.) (4,72) and water stress reduced
extension growth, node number and leaf area (138). Ten and 25
ml water/day/coffee seedling in 15 cm pots resulted in
drastic growth reduction, wilting and necrosis of the
terminal shoots compared to a 200 ml treatment which caused
water logging and induced leaf abscission (72).
Water stress has been shown to cause specific changes in
plant carbohydrate status. Decreased starch concentration
simultaneous with increased sugar content during drought
stress have been reported for sugar maple (Acer saccharum
Marsh.) (106) and black oak (Quercus velutina Lam.) (107). In
sugar maple total carbohydrates were not affected, indicating


11
a change in carbohydrate partitioning. Similar changes in
carbohydrates occurred in the inner bark of loblolly pine
(Pinus taeda L.) (56) and in cotton shoots (36). Water stress
induced increases in sugar concentration may serve several
purposes, of which two are of major significance. Increased
sugars increase the readily available substrate for
respiration (17). A second effect of increased free sugars
during water stress is to lower the osmotic potential of the
cell solution and hence maintain cell turgor. Concentration
and form of carbohydrates, organic acids or inorganic ions
are modified to promote osmotic regulation (74). A more
negative leaf solute potential with water stress was observed
in seedlings of English oak (Quercus robur L.) and silver
birch (Betula pendula Rodt.) but the degree of osmotic
regulation varied between species (105).
Munns and Pearson (98) found that reduced photosynthesis
as a result of water stress was not caused by accumulation of
photosynthates in potato leaves. Low leaf water potential
resulted in decreased translocation of carbohydrate, which
was proportional to the decline in net photosynthesis,
irrespective of whether tubers were present. A lower
proportion of labelled 1^C02 was found in tubers of stressed
potato plants compared to non-stressed plants (125).
One of the earliest plant responses to water stress is
reduced elongation of leaves, stems and roots (57).
Boyer (13) found that leaf enlargement of corn and soybean


12
was inhibited earlier and more severely than photosynthesis
and respiration by decreasing water potentials.
Physiological Effects of Root-Zone Temperature
According to Levitt (85) the two major forms of stress
injury are direct and indirect injury. Direct heat injury
results from a short exposure to an extreme temperature and
is detectable immediately at the cellular level. Indirect
injury results from prolonged exposure to temperatures below
those causing direct injury.
Supraoptimal temperature stress may be defined as the
retardation or cessation of metabolic functions in response
to high temperatures. Alexandrov (3) reported that the first
symptom of high temperature was the cessation of protoplasmic
streaming. This was followed by a reduction in photosynthesis
with subsequent damage to the chioroplasts. Finally,
semipermeabi1ity of cell membranes was disrupted, so that
cellular compartmenta1ization was lost with mixing of
cellular contents.
Starvation, a common form of indirect heat injury in
plants, occurs because of a higher optimum temperature for
respiration than photosynthesis (85). The temperature at
which photosynthesis and respiration rates are equal is the
temperature compensation point. When plant temperatures
exceed this critical point, carbohydrate reserves become
depleted and starvation ensues. Other forms of indirect
metabolic injury (85) are toxicities resulting from


13
disturbance of specific metabolic processes and biochemical
lesions resulting from the accumulation of metabolites
necessary for plant growth.
Ingram and Buchanan (61,62) demonstrated that electrol
yte leakage through root cell membranes in response to
increasing supraoptimal temperatures was sigmoidal and the
midpoint of this response was defined as the critical
temperature. Critical temperature was species and cultivar
specific and varied from 48 to 53C for a 20 minute exposure
time (61,62). Direct injury at high temperature was also
shown to be a function of exposure time (60).
Proteins are denatured by high temperature (2) and there
is evidence that high temperature disrupts membrane
structure. Inhibition of photosynthesis at high temperature
was correlated with the disruption of membrane associated-
processes including photosystem II (108),
photophosphorylation (123) and the change of chlorophyll
organization (122). The stability of membranes is mainly due
to the degree of fatty acid saturation which is linked to
membrane function and organelle association (145).
Environmental temperatures have been demonstrated to alter
fatty acid saturation ratios and membrane fluidity as
adaptive responses to high and low temperatures
(24,27,28,45,87).
There are several reports of the effects of growth
medium temperature on water uptake by roots (70,89,104,115).
Water conductivity through roots appears to be a temperature-


14
dependent process (117). Low temperature reduced
transpiration and stomatal conductance in chi 11-susceptible
species by reducing conductance through root cell membranes
(75,104,147). Such results are most likely due to increased
resistance to water at the endodermis layer (89).
Heating the root-zone to temperatures within the optimum
range generally improved plant growth (12,42,83) and heating
the root medium to 20C improved foliage plant production at
low greenhouse air temperatures (12). Root temperatures above
or below optimum may alter hydraulic conductivity and thus
indirectly affect stomatal conductance, photosynthesis and
subsequent carbohydrate translocation (66). Optimal root-zone
temperature in tomato (Lycopers icon esculentum Mill.) ranged
from 30 to 36C for nutrient translocation and water
absorption (42,58).
Using plants with heat-killed roots, Kramer (73) showed
that plant shoots remained alive and did not wilt for several
days even after root death. Transpirati on decreased after
root death because of leaf injury and gum deposits from dead
cells. Citrus hydraulic root conductance increased linearly
with increasing root temperature (137) and Carrizo citrange
exhibited a log-linear decrease in conductivity over a
temperature range from 40 to 10C (146).
Cooper (30) published an extensive literature review in
1973 on RZT effects and reported important differences due
not only to species and cultivars but also to growing
conditions. High leaf photosynthetic rates in soybean grown
r


15
at increasing RZT were related to low stomatal resistance,
high transpiration rate and high phosphoenol pyruvate and
RUDP carboxylase content (35). Cucumber (Cucumis sativus L.)
leaves were larger, contained more chlorophyll and produced
greater amounts of ATP and NAD PH when plants were grown at
25C RZT compared to 15 or 35C (26). Canopy photosynthesis
of pepper plants was closely related to shoot dry weight and
crop yield (44), although maximum growth and yields were
obtained at 30C RZT and maximum photosynthesis rates were
recorded at 36C RZT. The disparity between optimal RZTs was
explained by increased rates of respiration at the higher
RZT. High rates of photosynthesis at high RZT in tomato were
due not only to increased plant size but also to a
modification of the physiological and morphological propert
ies of the photosynthetic apparatus (43). Chlorophyll content
(26), enzyme activity (35) and leaf hormonal content (91)
have been influenced by increasing RZT.
Although many physiological processes are influenced by
high RZTs (30), there is relatively little information on the
effects of RZT on endogenous hormone levels (76). High RZTs
have been reported to increase gibberellin and decrease
cytokinin translocated in the root exudates (47,64,129).
There is also evidence that high RZT may affect the
distribution of gibberellin in the shoot (76). Alternatively,
high RZT may have increased the activity in the roots of
gibberellins previously synthesized in the shoot as well as
influenced synthesis in the shoot (110).


16
In a study on flowering in coffee (92), ambient
temperatures above 33C induced formation of orthotropic
shoots and prevented flower initiation. Root-zone heating
delayed flowering and induced excessive vegetative growth in
peppers (44). Such an antagonism between vegetative and
reproductive growth prompted the authors to recommend a
reduction in night temperature or nitrogen fertilization when
pepper plants are grown at high RZTs.
Reduced biosynthesis of cytokinins induced by supraopti-
mal RZT was thought to reduce photosynthetic rates in bean
(Phaseolus vulgaris L.) (21) and grape (Vitis vi nifera L.)
(129). Increasing RZTs from 18 to 30C in rice (Oryza sativa
L.) increased the translocation and rate of
photosynthetically assimilated l^C into the roots but
decreased root growth apparently by retarding protein and
cell wall synthesis (96).
Temperature has long been recognized as the main
environmental determinant of plant growth and phenology (81).
Increased soil temperatures have generally increased shoot
growth curvi1inearly and reduced root growth (30,112). All
growth variables of Pittosporum tobira Thunb. were
substantially lowered by a 40C RZT for 6 hrs/day compared to
27C (65), and Ingram (59) noted marked growth inhibition of
woody plants stressed by 35 to 40C RZT for 6 hrs/day.
Gosselin and Trudel (44) reported maximum leaf area in
pepper at a RZT of 30C with a decline at higher
temperatures. Watts (144) demonstrated reduced leaf expansion


17
in maize with increasing RZT but could not explain this as a
result of increased plant water deficits. RZTs above 37C
restricted leaf growth and rate of emergence in pearl millet
(Pennisetum typhoides S&H) (103). Soil temperatures of 37C
and above, which are common in the tropics, reduced both root
and shoot growth of maize and cowpea (Vigna unguiculata L.)
(90). Well-watered potted eggplants (Sol an urn melongena L.)
exhibited decreased plant growth with increasing RZT from 25
to 40C (118). Shoot to root ratio, however, was constant
except at 40C, where root rot occurred.
Philpotts (109) reported a linear decrease in cowpea
nodulation and total plant dry weight with increasing RZT
from 31 to 40C. RZTs above 32C significantly reduced
vegetative growth of two cultivars of cowpea through their
effects on shoot, peduncle and root dry weight and, to a
lesser extent, leaf number (94).
Using a system of circulating water coils, Franco
(39) applied RZTs from 13 to 48C and observed the effects
on coffee growth, transpiration and mineral absorption.
Maximum transpiration occurred at 33C with a significant
decline at 43C. Similarly, RZTs above 33C reduced
absorption of several nutrients and induced leaf chlorosis.
Plants growing at 48C died and both shoot and root growth
were depressed above 33C. In another experiment using young
coffee seedlings grown at 33C RZT, he reported the
occurrence of small tumors at the base of the stem from which
new orthotropic or non-flowering shoots grew.


18
In an extensive investigation using 'William' banana
in sunlit growth chambers, Turner and Lahav (142) reported
heat injury at an air temperature of 37C. Total plant weight
was greatest at 28C while leaf area was maximum at 33C.
Temperature altered partitioning patterns in the whole plant
and at 33C more leaves and less stem and roots were
produced, giving increased shoot to root ratios. Lamina leaf
folding also occurred at high air temperatures and the
authors implied that induced high vapor pressure deficits may
have been the cause. Severe constraints on data measurements
were imposed by the large nature of the banana plants and
restricted growth chamber space. This could, in part, explain
why critical physiological data such as photosynthesis,
transpiration and leaf water potential were not recorded.
Assimilate partitioning results were estimated using relative
growth rate formulae and no analyses on tissue carbohydrate
status were performed.
Root-Zone Temperature and Soil Moisture Interactions
The complexity of responses of RZT and soil moisture
stress alone are such that few researchers have attempted
studies on their interactive effects. Lai (80) reported in
1974 that a RZT of 35C significantly decreased root and
shoot growth and transpiration in maize seedlings. Effects of
high RZT were aggravated by low soil moisture levels. Changes
in soil water supply and RZT were found to have predictable


19
effects on the relationship between leaf water potential and
transpiration rate in several tree species (37,54,68,135).
Kaufmann (69) evaluated the effects of RZT (6 to 20C)
and two soil drying cycles (9 and 12 day) on water relations
and growth of Monterey pine (Pinus radiata. Don) under
greenhouse conditions. The slope of the relationship of xylem
pressure potential to transpiration was influenced by RZT.
Under well-watered conditions, neither transpiration nor
stomatal conductance were affected by RZT because xylem
pressure potential was not low enough to cause stomatal
closure. At the end of 9- and 12-day drying cycles, however,
transpiration was lower than in the well-watered controls as
a result of stomatal closure. RZT apparently had no consist
ent effect on transpiration rate. Reduced RZTs and soil
drying both significantly reduced shoot and root extension
after 28 days even though low RZT did not cause stomatal
closure.
In an earlier study with 'Monterey' pine, Babalola et
al. (5) measured photosynthesis, transpiration and
respiration at four soil water levels and four RZTs (10 to
27C ). They found that the rates of photosynthesis, respi
ration and transpiration decreased with increasing soil water
deficits. However, only at soil water levels lower than -0.7
bars and at RZTs of 10C and above were the rates of
photosynthesis and transpiration affected similar to changes
in stomatal conductance.


20
Barlow et al. (7) studied the effects of RZT and soil
water potential on corn seedlings. This investigation also
did not involve supraoptima1 temperatures but there were
notable interactive effects. Rates of leaf elongation, net
photosynthesis, transpiration and leaf water potential were
simultaneously monitored under varying stress conditions. In
their first study, leaf elongation was found to be much more
sensitive to changes in root temperature (12 to 28C) than
either photosynthesis or transpiration and ceased at a LWP of
-0.9 MPa; physiological responses were not affected until
LWPs of -1.2 to -1.3 MPa were attained. A decrease in net
photosynthesis coincided with an increase in stomatal and
mesophyll resistance at low LWPs. The authors concluded by
emphasizing the need for whole plant studies so that stress
effects on photosynthesis, translocation and assimilate
partitioning could be better analyzed.
Two root temperatures (21.1 and 26.6C) and two soil
water potentials (-0.35 and -2.50 bars) were employed in
their second experiment (8). Leaf elongation and total dry
matter were decreased 44% and 26%, respectively, soluble
carbohydrates increased 42% while rates of transpiration
decreased 24% at the low soil water potential. There were
also some interactions between RZT and soil water content.
Dry weights for plants at 21.1C and -0.35 bars were similar
to those at 26.6C and -2.50 bars. Higher soluble
carbohydrate concentrations with increased water stress were
offset by increased RZT. Decreased total plant dry weight and


21
transpiration at lower soil water levels were slightly offset
by higher RZT, presumably through increased root conductance
and water absorption.
Most of the studies involving soil temperature and
moisture interactions on banana in the tropics have been
field oriented and have involved the use of soil mulches
which effectively reduce soil temperature and conserve soil
moisture (10,11). In most cases, mulching led to increased
WUE which was based on a yield to evapotranspirati on ratio
rather than on the physiological parameters of photosynthesis
and transpiration. In the case of coffee, there are
independent studies on soil temperature (39) and soil
moisture (4,72,138), but no reports on their interactive
effects


CHAPTER III
EFFECTS OF IRRIGATION VOLUME ON BANANA AND IXORA UNDER
TWO GROWING CONDITIONS
Introduction
Although 'Grande Naine' banana (Musa spp. AAA) is a
mesophytic, herbaceous plant requiring moist soils for
optimum production (127), it can be grown in marginal areas
where soil water is limiting. Irrigation scheduled when
available soil moisture depletion reaches 60% to 80% has been
shown to be optimum (10,11,31,116), but detailed
characterization of physiological responses in banana to
decreasing soil water levels is lacking.
Moisture stress is thought to be necessary for flower
induction (4) in coffee, a close relative of ixora (Ixora
chinensis L. 'Maui') within the Rubiaceae family (6), but
adequate irrigation (72) is essential for growth and mineral
absorption (138). Because of its importance as a major
beverage crop, there are several field (100,101) and
container studies (39,72,138,140) on the physiological
responses of coffee, but no work has been conducted on the
effects of varying soil water levels on container-grown
ixora. The present study investigated the physiological
responses of banana and ixora to three irrigation levels and
22


23
a 14-day drying cycle. Irrigation experiments were conducted
under both greenhouse and growth room conditions.
Materials and Methods
Plant Materials and General Cultural Procedures
Ten- to 12-cm tall tissue-cultured banana plants and
similar-sized, uniform ixora rooted cuttings were obtained
from commercial nurseries. Plants were hardened with
intermittent mist (6 sec min-1) in a glass greenhouse at 28C
under 80% light exclusion for one week, then moved to 40%
light exclusion for another week. Plants were transplanted to
white 4-cm diameter x 21-cm tall conical containers (150 cm3)
using Metro-Mix 300 growth medium (W.R. Grace and Co.,
Cambridge, MA). Plants were then moved to the experimental
greenhouse or growth room where they were watered to
container capacity daily and allowed to acclimatize for one
week prior to the initiation of the experiment.
Greenhouse
Experiment 1. Experiment 1 was initiated in July, 1985,
in an air-conditioned greenhouse in which maximum
photosynthetic photon flux density (PPFD) ranged from 700 to
800 umol m-2 s-1, and temperatures were 25 to 30C day and 18
to 21C night. Relative humidity varied from 40 to 80%
(Figure 3-1).
Plants were watered daily at 2200 hr with 10+2 ml (Wl),
20+4 ml (W2) or 40 + 8 ml (W3) per container. These irrigation


PPFD (pmol
0800 1000 1200 1400 1600
TIME (hrs)
Figure 3-1. Photosynthetic photon flux density and relative humidity in the
greenhouse during measurements of physiological parameters.
I\>
-P*
PERCENT RELflTIUE HUniDlTYO


25
volumes provided 53% to 60%, 65% to 75%, and 85% to 100%
container capacity (CC) respectively, for banana and 52% to
64%, 68% to 80% and 90% to 100% CC, respectively, for ixora.
The comparatively smaller root mass of ixora was responsible
for the difference in growth medium waterholding capacities
between these plants. Water was applied automatically through
drip tubes by a battery-operated controller (Water Watch
Corp. Seattle, WA). One, two and four 1-mm drip
tubes/container constituted the Wl, W2 and W3 irrigation
treatments, respectively.
After 14 days of irrigation treatments, diurnal
measurements of leaf photosynthesis (PS), transpiration (TR)
and leaf conductance (CS) were made using a portable
photosynthesis system (Model LI-6000, LI-C0R, Inc., Lincoln,
NE). Measurements were initiated at 0800 hr EDT on a
cloudless day and taken every 2 hr until 1600 hr. A 1-liter
cuvette chamber was used for measurements and the mean of
eight consecutive 30-sec observations on each leaf
constituted a measurement. A zero check of the analyzer was
performed between treatments within each replicate.
Simultaneous measurements of LWP were made using a Scholander
type pressure chamber (PMS Instrument Model 600, Santa
Barbara, CA) as described by Barrett and Nell (9). The third
most recently expanded leaf in banana was selected for
measurements of both gas exchange and LWP. This leaf has been
shown to be the most responsive and the youngest with fully
developed stomata (127). Gas exchange processes in ixora were


26
measured on the most recently matured leaves but apical
shoots were sampled for LWP. Six plants per treatment were
sampled for all measurements.
Experimental design was a randomized complete block with
21 plants per water treatment within each of three blocks.
Six replicate plants were randomly selected at each sampling
time. Water use efficiency (WUE) was calculated from PS/TR.
Means and standard errors were calculated and plotted against
time of day.
Experiment 2. In order to further monitor physiological
responses of water stress and correlate these with visual
plant symptoms, water was withheld from banana and ixora
plants for 14 days beginning one week after transplanting.
This drying cycle study was conducted simultaneously with
experiment 1 in the same greenhouse and similar cultural
practices were used. Midday LWP and CS measurements were
recorded on five plants of each species at 2-day intervals
using the procedures described in experiment 1. A completely
randomized design was used in this experiment and means and
standard errors were calculated. There were five replicate
plants per treatment for each sampling date.
Growth room.
Experiment 3. Experiment 1 was repeated in a 3.0 m by
7.6 m walk-in growth room in order to investigate
physiological responses to water stress under more precisely
controlled environmental conditions. Irradiance of 1100 umol
m2 s-1 PPFD, as measured by a quantum radiometer (LI-COR


27
Model LI-185A, LI-COR Inc., Lincoln, NE), was supplied at
plant canopy level by 1000 W phosphor-coated metal-arc HID
bulbs (GTE Sylvania Corp., Manchester NH) from 0600 to 1830
hr daily. Air temperatures of 28C day and 21C night and a
relative humidity of 65% to 70% were maintained. Three
irrigation volume treatments were applied as in experiment 1
in a randomized complete block design with three blocks and
21 plants per water treatment per block. Species were treated
and analyzed as separate experiments. On the fifteenth day
after water stress treatments, diurnal measurements were
initiated at 0800 hr and continued at 2-hr intervals until
1800 hr. Leaf sampling methods and measurement procedures for
PS, CS, TR and LWP were employed as in experiment 1.
Results and Discussion
Greenhouse
Experiment l--banana. Irrigation treatments influenced
daily PS, CS and TR maxima as well as diurnal patterns
(Figure 3-2A, 3-2B, 3-2C). Relatively high LWP were
maintained by plants in the W2 and W3 irrigation treatments
but there was a distinct decline in LWP by 1000 hr in plants
receiving the lowest irrigation volume, W1 (Figure 3-2D).
PS patterns of plants under the W3 and W2 irrigation
treatments paralleled the corresponding CS patterns, with a
rise in PS from 0800 hr to a peak at 1400 hr of 0.20 mg CO2
m2 s-1 for the W3 treated plants. Maximum PS for plants in
the W2 irrigation treatment were reduced almost three-fold


Figure 3-2. Effects of three irrigation volumes on the diurnal physiological
responses of 'Grande Naine' banana grown under greenhouse conditions. A. leaf
photosynthesis B. leaf conductance C. transpirati on and D. leaf water potential.
W1 = 10+2 ml, W2 = 20+4 ml and W3 = 40 +_8 ml daily per 150 cm3 container. Points
are the means of six replicate plants and vertical bars represent the SE.


lTION (mg H20 m-2 s-1) PHOTOSYNTHESIS (mg 002 m-2
0


30
below plants at W3 (Figure 3-2A). PS for the W1 treated
plants was significantly reduced compared to plants in the
other irrigation treatments with a midday rate of only 0.003
mg CO2 m-2 s--*-.
CS in plants receiving the highest irrigation volume,
W3, averaged 0.37 cm s1 at 0800 hr, had more than doubled by
1400 hr and declined sharply to 0.17 cm s-* at 1600 hr
(Figure 3-2B). CS in plants under the W2 irrigation followed
the same pattern, but was reduced by 50%, 47% and 49% at
1000, 1200 and 1400 hr, respectively. CS in plants at the
lowest irrigation level did not change appreciably throughout
the day and averaged only 0.08 cm sl, suggesting that the
stomata remained nearly closed throughout the day.
Therefore, PS and CS were severely reduced by decreased
irrigation and both declined in parallel.
Although TR increased from 0800 hr to 1000 hr for both
W2 and W3 treated plants, TR in W2 and W1 plants was
significantly lower (Figure 3-2C). TR remained somewhat
stable between 1000 and 1400 hr for all treatments and
declined significantly between 1400 and 1600 hr under the W3
irrigation volume. Maximum TR was 44.4, 14.2 and 6.3 mg H2O
m-2 s-1 for the W3, W2 and W1 treated plants, respectively.
LWP of plants under the W2 and W3 irrigation treatments
were fairly stable throughout the day, but LWP of plants at
the lowest irrigation level declined significantly after 0800
hr to a low LWP of -0.90 MPa at 1000 and 1200 hr (Figure 3-
2D). Although there was an increase in LWP for W1 treated


31
plants after midday, the LWP at 1600 hr of -0.62 MPa remained
significantly lower than the 0800 hr LWP. The reduction in CS
and the decline in gas exchange processes but not leaf water
status at midday and 1400 hr for the W2 treated plants,
indicated that there were factors other than leaf water
status controlling stomatal opening.
Water use efficiency (WUE) for plants under the W3 and
W2 irrigation levels increased almost linearly throughout the
day but did not differ between treatments until 1200 hr,
after which WUE in W2 treated plants was higher (Figure 3-3).
This indicated that TR in plants under the W2 treatment was
reduced more relative to PS. However, the WUE pattern for the
severely stressed W1 treated plants remained low throughout
the day. The leaf folding mechanism commonly observed in
field-grown banana plants at midday hours (127,136) was
observed in the W2 treated plants. Increased WUE values,
though calculated on a yield/evapotranspiration ratio, have
also been obtained for field-grown bananas under moderate
reductions in soil moisture levels (11).
Altered physiological processes in banana with
decreasing irrigation levels agrees with Shmueli's work (126)
in which TR and CS in Cavendish banana were reduced by
available soil moisture levels below 66%. Chen (25) also
found a similar effect at a soil water level of 50% to 60%
field capacity. These were both field studies in which soil
moisture effects could have been confounded by ambient vapor


Figure 3-3. Effects of three irrigation volumes on the diurnal water use
efficiency of 'Grande Naine' banana grown under greenhouse conditions. W1
ml, W2 = 20+4 ml and W3 = 40jj8 ml daily per 150 cm^ container. Points are
means of six replicate plants and vertical bars represent the SE.
= 10 + 2
the


0800 1000 1200 1400 1600
UflTER USE EFFICIENCY (mg C02/mg H2O.J0-4)
cJ en (_o
q Q Q Q
ee
120


34
pressure deficits to a greater extent than occurred in the
present greenhouse experiment. CS and TR reported for both
field studies, however, were closely correlated. Brun (20)
reported good correlation between PS and TR in intact banana
leaves in response to light intensity but CS was not
monitored in his study. Results in the present experiment
indicate that stomatal closure was associated with reductions
in PS and TR.
Severe growth reduction and leaf chlorosis have been
induced by soil moisture depletion levels below 50% in field
studies (126). These effects were generally attributed to
reductions in PS, although PS rates were not measured. The
present study substantiates these hypotheses, since the
lowest irrigation level produced similar water stress
symptoms and significantly decreased PS (Figure 3-2A).
Experiment l--ixora. Although irrigation treatments
highly influenced measured parameters, diurnal patterns did
not necessarily parallel one another (Figure 3-4). Under the
W3 irrigation treatment, maximum PS and TR rates of 0.26 mg
CO2 m-2 s"1 and 58.1 mg H2O m-2 s-*, respectively, occurred
at 1400 hr (Figure 3-4A, 3-4C). However, CS was somewhat
erratic, with a maximum at 1000 hr, followed by a decline at
1200 hr and a small rise at 1400 hr (Figure 3-4B). PS and CS
were significantly reduced by the W2 and W1 irrigation levels
through 1400 hr compared to W3 and under the W1 level CS and
TR appeared to be more closely related.


Figure 3-4. Effects of three irrigation volumes on the diurnal physiological
responses of Ixora chinensis L. 'Maui' grown under greenhouse conditions. A. leaf
photosynthesis 6. leaf conductance C. transpiration and D. leaf water potential.
W1 = 10+_2 ml, W2 = 20+4 ml and W3 = 40+8 ml daily per 150 cm3 container. Points
are the means of six replicate plants and vertical bars represent the SE.


0800 1000 1200 1400 1600 ^aO 1000 1200
TIME (hrs) TIME (hr)
TRANSPIRATION (mg H20 m-2 s-1) PHOTOSYNTHESIS (mg 002 m-2 *-1)
9e


37
LWP was generally reduced by decreasing irrigation
volumes (Figure 3-4D). Midday LWP for the W2 and W1
treatments were 2 and 2.5 times less than for plants under
the W3 irrigation treatment, respectively. Irrigation
treatments, therefore, were well distinguished by measuring
LWP.
Reduced irrigation, however, decreased PS relatively
more than TR as reflected in the higher WUE in plants at the
W3 irrigation level compared to plants under the W2 and W1
levels (Figure 3-5). Non-stoma tal inhibition of PS under
water stress conditions have also been reported in other
species (14,15) and may have contributed to the lower WUE
observed with water stress in this study.
Although midday declines in CS have been noted for
coffee (100,101,140) under well-watered field conditions, the
suggestion that light was responsible (101) would not seem to
apply to ixora. In the high light environment of the growth
room CS did not decline at midday (Figure 3-10B).
A high irrigation treatment of 200 ml/15 cm pot/day
induced flooding effects and caused leaf abscission in coffee
(72). Such effects were not induced in this study by the
highest irrigation treatment, W3, but the erratic midday
stomatal response referred to above might have been in
response to the high watering level.


Figure 3-5. Effects of three irrigation volumes on the diurnal water use
efficiency of Ixora chinensis L. tMaui grown under greenhouse conditions. W1
10 + 2 ml, W2 = 20 + 4 ml and W3 = 40 + 8 ml daily per 150 cm^ container. Points are
the" means of si x~repl i cate plantsand vertical bars represent S.E.


UIRTER USE EFFICIENCY (mg C02/,,n9
&
&
J
O
o
CM
x
60
45
15
0
Ull
U2
0800
1000 1200
1400
1600
TIDE (hrs)
co
co


40
Experiment 2--banana. LWP decreased linearly while CS
decreased in a sigmoidal pattern during the 14 days without
irrigation (Figure 3-6). Leaf folding along the midrib is a
common feature of water-stressed field-grown banana plants at
midday hours (127,136), but plant water status associated
with the mechanism has not been previously defined. In this
study, leaf folding occurred between day 4 and 6 and
coincided with a decline in LWP from -0.51 to -0.65 MPa and a
decrease in CS from 0.46 cm s"l on day 4 to 0.051 cm s"1 on
day 6. In experiment 1, leaf folding was also observed in the
W2 treated plants at midday with comparable LWP and CS. This
response therefore, substantiated the irrigation-volume
method as a fairly accurate, practical means of imposing
water stress treatments in banana. Anatomical studies have
indicated a special band of cells along the midrib of the
banana leaf (130) which are responsible for the folding
response and it is likely that these cells are more
responsive to decreasing LWP than the other leaf cells.
Further reduction in LWP to -0.91 MPa and CS to 0.019 cm
s_1 probably impaired the PS mechanism since chlorosis
occurred indicating greater chlorophyll degradation than
synthesis (57). At day 14, when LWP declined to -1.34 MPa and
CS to 0.004 cm s"l, there was severe chlorosis and apparent
growth cessation. At this stage leaves remained folded
probably indicating a cessation in gas exchange processes and
complete stomatal closure.


Figure 3-6. Effect of a 14 day drying cycle on leaf water potential and leaf
conductance of 'Grande Naine' banana grown under greenhouse conditions. Points
are the means of five replicate plants and vertical bars represent the SE.


42
Experiment 2--ixora. LWP decreased linearly after day 2
but there was a steep decline in CS from 0.53 cm s_1 on day 2
to 0.14 cm sl on day 4 (Figure 3-7). Wilting of youngest
leaves occurred between day 4 and 6 and corresponded to
decreases in LWP from -1.08 to -1.71 MPa and in CS from 0.14
to 0.038 cm s"1. Severe wilting of top leaves and abscission
of bottom leaves were observed on day 8 at a LWP of -2.06 MPa
and a CS of 0.008 cm s~l. Plant water status between days 4
and 6 of the cycle was comparable to that at midday
(-1.2 MPa) in plants under the W1 irrigation treatment in
experiment 1. This indicated that reducing irrigation volume
was an effective means of imposing water stress in ixora.
Beyond the day 8 stage, wilting and leaf abscision made it
difficult to record further data. Since stressed plants were
not rewatered it was impossible to determine whether stomatal
closure and leaf abscission observed in this study were
clearly mechanisms imparting stress tolerance that allowed
the plants to survive.
Growth Room
Experiment 3--banana. PS, CS and TR diurnal patterns
paralleled one another under the more controlled conditions
of the growth room (Figure 3-8) but irrigation treatment
effects were not quite as dramatic as in the greenhouse
experiment (Figure 3-2). Maximum PS, CS and TR occurred in
the growth room at 1000 hr followed by a gradual decline to


Figure 3-7. Effects of a 14 day drying cycle on leaf water potential and leaf
conductance of Ixora chinensis L. 'Maui' grown under greenhouse conditions.
Points are the means of five replicate plants and vertical bars represent the SE.


Figure 3-8. Effects of three irrigation volumes on the diurnal physiological
responses of 'Grande Naine' banana grown under growth room conditions. A. leaf
photosynthesis B. leaf conductance C. transpiration and D. leaf water potential.
W1 = 10+2 ml, W2 = 20+4 ml and W3 = 40+8 ml daily per 150 cm^ container. Points
are the means of six replicate plants and vertical bars represent the SE.


TRANSPIRATION (mg H20 nrr-2 8-1) PHOTOSYNTHESIS (mg C02 m-2 s-2]


46
1800 hr (Figure 3-8) while maximum levels in the greenhouse
experiment were at 1200 to 1400 hr for CS and 1000 to 1400 hr
for TR. Higher light intensity early in the day in the growth
room could have resulted in earlier stomatal opening (33).
Both Shmueli (126) and Chen (25) reported CS and TR diurnal
maxima in banana at 1000 hr followed by declines and with
minor increases at 1400 hr.
Under the W3 irrigation treatment, maximum PS, CS and TR
were 0.596 mg CO2 m~2 s"*, 0.720 cm s* and 89.2
mg H2O m-2 s-1, respectively at 1000 hr. PS, CS and TR were
reduced 252, 482 and 352 at 1000 hr, respectively, by the W2
irrigation treatment compared to the W3 treatment (Figure 3-
8A, 3-8B, 3-8C). At the lowest irrigation level, all gas
exchange processes remained consistently low throughout the
day and maximum PS, CS and TR were 0.097 mg CO2 m-2 s"*,
0.181 cm s"1, and 38.2 mg H2O m-2 s*, respectively.
LWP generally declined for all irrigation treatments
after 0800 hr for most of the day with an increase after 1400
hr for the W3 treated plants (Figure 3-8D). Midday LWPs were
-0.58 MPa, -0.66 MPa and -0.82 MPa for the W3, W2 and W1
treatments, respectively. LWP between 1000 hr and 1400 hr
were not different for the W3 and W2 treatments but LWP was
significantly reduced by the W1 irrigation level. Despite the
similar LWP between W2 and W3 at 1000 to 1400 hr, significant
differences in PS, CS and TR were observed.
Davies (33) reported increased stomatal sensitivity to
decreasing soil water levels for cotton and soybean under


47
growth chamber conditions but Boyer (14) found PS of
sunflower to be affected more by photochemical activity than
by stomatal conductance under similar conditions. In the
present study there were relative increases in PS but not CS
in plants grown in the growth room as compared to the
greenhouse.
Maximum CS and TR have been shown to occur in moderately
watered field-grown banana at 1000 hr with subsequent
declines and smaller peaks at 1400 hr (25,126). It is notable
that this response resembled the diurnal patterns that were
observed in the W2 treated plants in the present study. In
the growth room LWP were generally lower than in the
greenhouse and could have been responsible for the general
decline in PS, CS and TR after 1000 hr.
Under growth room conditions, the W3 irrigation
treatment generally maintained a constant WUE throughout the
day (Figure 3-9) reflecting the parallel changes in PS and
TR. Though WUE for the W2 treated plants declined at 1200 and
1400 hr compared to 0800 hr, it was still not significantly
different from that of the W3 plants during these hours. The
relative improvement in WUE at the lowest irrigation in this
experiment compared to the same treatment in the greenhouse
study was due mainly to increased PS. The W1 irrigation
treatment under growth room conditions therefore did not
depress physiological responses as much as that observed in
the greenhouse study.


Figure 3-9. Effects of three irrigation volumes on the diurnal water use
efficiency of 'Grande Naine' banana grown under growth room conditions. W1 = 10+_2
ml, W2 = 20+4 ml and W3 = 40+8 ml daily per 150 cm^ container. Points are the
means of six replicate plants and vertical bars represent the SE.


I
o
o
(N
X
cn
E
\
(N
O
U
cn
E
U
z
u
LJ
Ll)
in
3
CL
CL
n
120 -
80 --
40 --
0
0800
1000
1200 1400 1 BOO 1800
TinE (hrsJ
-p*


50
Experiment 3--ixora. Linder growth room conditions, the
W2 and W1 treatments did not consistently decrease
physiological parameters (Figure 3-10) as severely as in the
greenhouse study (Figure 3-4). At 1200 hr, there were no
differences in responses of plants under the W2 and W3
treatments but at the lowest irrigation treatment, all
parameters were significantly reduced over those at the
higher irrigation levels.
Maximum PS rates were 0.425, 0.388 and 0.151 mg CO2 m~2
s"l for the W3, W2 and W1 irrigation treatments, respectively
(Figure 3-10A). These rates were 1.25, 7 and 10 times higher
than the respective rates in the greenhouse study and may
have reflected the influence of increased PPFD on PS in
ixora, particularly at the lower irrigation treatments. Light
levels in the greenhouse were probably below light saturation
levels for ixora.
There were no differences in CS between plants at the W2
and W3 irrigation levels from 0800 to 1400 hr but CS was
significantly reduced at these hours by the W1 irrigation
volume (Figure 3-10B). Midday CSs were 0.492, 0.382 and 0.190
cm s-1 under the W3, W2 and W1 treatments, respectively.
Compared with CS in plants in the greenhouse experiment
(Figure 3-4B), these values were 342, 492 and 872 higher at
the W3, W2 and W1 irrigation levels, respectively. The
increased response in stomatal opening, especially at the
lower irrigation levels corresponded with increases in PS.


Figure 3-10. Effects of three irrigation volumes on the diurnal physiological
responses of Ixora chinensis L. 'Maui1 grown under growth room conditions. A.
leaf photosynthesis B. leaf conductance and C. transpiration. W1 = 10+2 ml, W2 =
20+4 ml and W3 = 40+8 ml daily per 150 cm^ container. Points are the means of six
replicate plants ancf vertical bars represent the SE.


PHOTOSYNTHES8 (mg 002 m-2 t-1)
p *.,* s a.


53
The high PS rate at 0800 hr under the W3 irrigation
resulted in a correspondingly high WUE (Figure 3-11) and
although WUE then declined at 1000 and 1200 hr for the
highest irrigation treatment, WUE values were basically the
same for all irrigation treatments through the rest of the
day. Under greenhouse conditions, W3 treated plants clearly
had higher WUE than plants subjected to lower irrigation
levels (Figure 3-5). The increase in WUE in the W2 and W3
relative to the W3 treated plants under growth room
conditions was due mainly to the increased PS rates in the
high light environment.
Growth room conditions, therefore, not only moderated
physiological responses of ixora to reduced irrigation volume
but also stabilized PS/TR changes as reflected by WUE values.
The increased photosynthetic response in plants grown under
the W1 and W2 irrigation volumes suggested that ixora could
probably better withstand decreased irrigation volume in the
growth room compared to greenhouse conditions reported in
this study.


Figure 3-11. Effects of three irrigation volumes on the diurnal water use
efficiency of Ixora chinensis L. 'Maui' grown under growth room conditions. W1
10+2 ml, W2 = 20 + 4 ml and W3 = 40+^8 ml daily per 150 cm3 container. Points are
the means of six replicate plants and vertical bars represent the SE.


UlflTER USE EFFICIENCY (mg C02/mg
i
o
o
(N
X
80
40
0
I 1 1 1 1 h-
0800 1000 1200 1400 1600 1800
TinE (hrs )
cn
tn


CHAPTER IV
ROOT-ZONE TEMPERATURE EFFECTS ON BANANA AND IXORA
UNDER TWO GROWING CONDITIONS
Introduction
High root-zone temperatures (RZT) have been shown to
reduce plant growth (59,63,70) and affect many physiological
processes (43,46,64). Tropical and subtropical plant species
are often thought to be heat tolerant, but soil temperatures
as high as 52C have been recorded in the tropics (39), and
temperatures in this range can be lethal for some tropical
(39,63) and subtropical crops (62). An air temperature of
33C was reported as being optimum for growth and dry weight
partitioning in banana (Musa spp. AAA) (142) and mineral
composition was highly influenced by temperatures from 18 to
33C (143). A RZT of 33C was found to be optimum for growth
and transpiration in coffee (39), a member of the same
subfamily of the Rubiaceae to which ixora belongs (6).
However, there are no reports on RZT effects on growth and
physiology of container-grown banana or ixora.
It is essential to identify growth and physiological
effects of high RZT in container production systems before
control measures can be developed to alleviate such effects.
This study investigated the short-term effects of RZT on
56


57
container-grown 'Grande Naine' banana and Ixora chinensis L.
'Maui' under greenhouse and growth room conditions.
Materials and Methods
Plant Materials and General Cultural Procedures
Ten- to 12-cm tissue-cultured 'Grande Naine' banana
plants and similar-sized, uniform I xora c hin e n sis L. 'Maui*
rooted cuttings were obtained from commercial nurseries.
Plants were hardened in intermittent mist (6 sec min"1) for
one week under 80% light exclusion and then moved to 40%
light exclusion for another week. Plants were then
transplanted to 4-cm diameter x 21-cm high conical containers
(150 cm^) using Metro-Mix 300 growth medium (W.R. Grace and
Co., Cambridge, MA). Plants were fertilized weekly with a
soluble 20N-8.8P-16.6K fertilizer (Peters 20-20-20, W.R.
Grace and Co., Cambridge, MA) at 150 ppm N.
Experiment 1--Greenhouse
An experiment was initiated August, 1985 in an air-
conditioned glass greenhouse with a mean maximum PPFD of 600
to 700 umol m"2 s-* PPFD, and a 25 to 30C day and 18 to
21C night temperature. Relative humidity was not controlled
and varied from 40% to 80% (Figure 4-1). Recently
transplanted banana and ixora plants were watered daily to
container capacity and allowed to acclimatize for one week.
RZT treatments were established within styrofoam-lined
wooden air bath boxes (1 m x 1 m x 20 cm) in which plant
containers were firmly inserted to within 2 cm from their


I ornr) Qjdd
100
O
*
1000
X
- 800
I
Ul
CN
i
E
600
400
200
0
80
60
40
20
0800 1000 1200 1400 1600
TjriE (hrs 1
Figure 4-1. Photosynthetic photon flux density and relative humidity in the
greenhouse during measurements of physiological parameters.
PERCENT RELBTHdE HUniDITY O


59
upper rims. Four equally spaced 100 watt aluminum foil-
covered incandescent light bulbs provided convective heating
of the enclosed root systems. Proper distribution of heat and
aeration within the boxes were ensured by two fans (IMC
Magnetics, Roch. NH) per box. The light bulbs in each box
were controlled by a pre-set thermostat and the temperatures
in each box were verified daily with a thermocouple
thermometer (TH 65, Wescor, Inc., Logan, UT). Electrical
power to the temperature boxes was controlled by Intermatic
Time Controls (Intermatic Inc. Spring Grove, Ill), providing
daily RZT treatments from 1000 to 1600 hr. Boxes were placed
on 1 m high benches and an automatic drip irrigation system
provided daily watering of all plants to container capacity
at 2200 hr.
RZT treatments established were 28^1, 33+_l, 38^1 and
43+_lC. Each temperature box contained six ixora and six
banana plants and there were three replicate boxes for each
treatment temperature arranged in a randomized complete block
design. Species were treated and analyzed as separate
experiments. Means and standard errors were calculated for
the six replicate plants measured at each sampling time.
After 14 days of RZT treatments, diurnal measurements of
leaf photosynthesis (PS), leaf conductance (CS),
transpiration (TR) and leaf water potential (LWP) were made
using a portable photosynthesis system (LI-6000 model, LI
COR, Inc., Lincoln, NE). Measurements were initiated at 0800
hr EDT on a cloudless day and taken every 2 hr until 1600 hr.


60
A 1-liter cuvette was used and the mean of eight consecutive
30-sec observations constituted a measurement. A zero check
of the analyzer was performed between treatments within each
replicate. Simultaneous measurements of LWP were made using a
pressure chamber (PMS Instrument Model 600, Santa Barbara,
CA) as described by Barrett and Nell (9). The third most
recently expanded leaf in banana was selected for both
gaseous exchange processes and LWP measurements. This leaf
has been shown to be the most responsive and the youngest
with fully developed stomata (127). The most recently matured
ixora leaves were used for PS, CS and TR measurements and 4-
to 5-cm shoot tips were sampled for LWP.
Experiment 2--Growth Room
In order to further monitor the physiological responses
of ixora and 'Grande Naine' banana to RZT under more
precisely controlled environmental conditions, experiment 1
was repeated in a 3.0-m by 7.6-m walk-in growth room. Light
irradiance of 1100 umol m-2 s-* PPFD, as measured by a
quantum radiometer (LI-C0R Model LI-185A, LI-COR Inc.,
Lincoln, NE), was supplied from 0600 to 1830 hr at plant
canopy height by 1000 W phosphor-coated metal-arc HID bulbs
(GTE Sylvania Corp., Manchester, NH). Air temperature of 28C
day and 21C night and a relative humidity of 65% to 70% were
maintained.
Each conical container with a transplant was suspended
through a tightly fitting styrofoam ring within a specially


61
constructed root heating tube (RHT). Each RHT was 22.5 cm
high and constructed from 7.5 cm diameter metal pipe wrapped
with 60 watt 120 vac heating tape (Smith-Gates Corp.,
Farmington, CT) and 1.25 cm thick foam insulation. The RHTs
were connected to solid state electronic controllers which
maintained preset treatment temperatures by a thermistor
feedback mechanism. Each controller maintained treatment
temperatures in 16 tubes with four tubes at each of four
specified temperatures. Treatment temperatures were set and
also maintained at 28+0.3, 33+0.3, 38+0.3 and 43+0.3C.
Similar leaf number and sampling methods were used and the
same physiological measurements were recorded as in
experiment 1 after 14 days of root-zone temperature
treatments.
Experimental design was a randomized complete block with
36 plants of each species per temperature treatment arranged
in three blocks. Species were treated and analyzed as
separate experiments. Means and standard errors were
calculated from six replicate plants measured at each
sampling time.
Results and Discussion
Experiment 1--Greenhouse
Banana. At the 33 and 28C RZTs, PS increased to midday
maxima of 0.53 and 0.50 mg C 0 2 m~^ $ 1 ^ respectively, and
then declined (Figure 4-2A). Maximum PS was 0.43 mg CO2 m2
sl at 1000 hr in the 43C RZT treated plants, while PS was


Figure 4-2. Effects of four root-zone temperatures on the diurnal physiological
responses of 'Grande Naine' banana grown under greenhouse conditions. A. leaf
photosynthesis B. leaf conductance C. transpiration and D. leaf water
potential. Points are the means of six replicate plants and vertical bars
represent the SE.


£9


64
0.31 mg CO2 m~2 S-1 f0r plants at the 38C RZT. These rates
were 202 and 422, respectively, of the maximum PS recorded at
33C.
Midday CS for the 28 and 33C RZT treated plants were
0.43 and 0.34 cm s*1, respectively, with significant
reductions to 0.28 and 0.19 cm s~l for plants at the 38 and
43C RZT (Figure 4-2B). TR diurnal patterns were closely
related to those for CS, with maximum rates of 54 and 52 mg
H2O m-2 s-1 at midday for plants at the 28 and 33C RZT,
respectively (Figure 4-2C). Midday TR was significantly
reduced by the 38 and 43C RZT compared to the other RZTs.
LWPs were generally not different throughout the day for
the 28, 33 and 38C RZT treated plants. Among these three
RZT treatments, a low midday LWP of -0.52 MPa was recorded at
the 38C RZT (Figure 4-2D). A significant reduction in LWP
was noted for plants at the 43C RZT at 1200 hr, with a
midday LWP of -0.76 MPa. Although transpirational water loss
was reduced in plants at the 43C RZT at midday compared to
plants at the 28 and 33C RZTs, LWP of plants at the highest
RZT were significantly lower than those of the other three
RZT treatments. At 38C, TR was less than at 28 and 33C RZT
but LWPs were the same between these treatments. These
responses may suggest impairment of the absorptive and/or
conductive capacity of roots in plants exposed to the 43C
RZT.
In summary, PS in banana plants was greatest at the 28
and 33C RZTs under greenhouse conditions. The 28, 33 and


65
38C RZT treatments resulted in the highest LWP and the 43C
RZT reduced all measured parameters. RZTs of 28 and 33C are
well within the range of RZTs reported as being optimum for
the growth of coffee (39), tomato (42) and pepper (44) while
38C RZT has reportedly decreased root conductance (146) and
transpiration (73).
Ixora. PS in plants at all RZT treatments increased from
0800 hr to midday (Figure 4-3) but PS, CS and TR were
greatest at midday in plants exposed to the 33C RZT. LWP
declined for most treatments at 1200 hr, with progressively
lower values from the 28 and 33C RZT to the 38 and 43C
treatments (Figure 4-3D).
Maximum midday PS rate was 0.41 mg CO2 m-2 s-1 for the
33C treated plants with reductions to 0.29, 0.28 and 0.25 mg
CO2 m-2 s"1 for the 38, 43 and 28C RZT treatments,
respectively (Figure 4-3A). At 1400 hr, however, PS rates at
the 43 and 38C RZT were 0.26 and 0.23 mg CO2 m-2 s*,
respectively, while PS at 33C RZT had declined to 0.17 mg
CO2 m-2 s-l. At 1600 hr, PS rates for the 28, 33 and 38C
RZT treated plants were not different. The 33C RZT clearly
resulted in the highest midday PS but because this RZT did
not alter morning rates and caused rapid post-midday
declines, mean daily PS at 33C was probably similar to that
at other RZT treatments.
CS and TR diurnal patterns were closely related, with
the midday maxima of 0.53 cm s-1 and 95 mg H2o m2 s_1,
respectively, for both parameters occurring in the 33C RZT


Figure 4-3. Effects of four root-zone temperatures on the diurnal physiological
responses of Ixora chinen sis L. 'Maui' grown under greenhouse conditions. A. leaf
photosynthesis BT leaf conductance C. transpirati on and D. leaf water
potential. Points are the means of six replicate plants and vertical bars
represent the SE.


0 0 0 0 1000 1200 1400 1600 08 0 0 1000 1200
TIME (hrs) TIME (hrs)
TRANSPRAT10N (mg H20 m-2 8-1)
o 8 § 8 §
i i
PHOTOSYNTHESIS (mg 002 m-2 8-1)
L 9


68
plants (Figure 4-3B, 4-3C). Midday CS and TR were
significantly reduced in plants at the 43C RZT compared to
the 33C RZT but the 38C RZT resulted in the lowest midday
TR and CS.
Mean midday LWP for plants at the 28 and 33C RZT was
-0.71 MPa with progressive reductions by the 38C (-0.87 MPa)
and 43C (-1.02 MPa) RZT treatments (Figure 4-3D).
The 33C RZT treated ixora plants exhibited highest
midday PS, CS, TR and had relatively good plant water status
compared with other RZT. Although PS was not measured, Franco
(39) reported that a RZT of 33C was optimum for coffee in
terms of TR and root and shoot dry weights. TR and plant dry
weights were reduced with increasing RZT above 33C and
plants died at 48C RZT.
Midday LWPs decreased significantly to -0.87 MPa by the
38C RZT treatment and to -1.02 MPa by the 43C RZT
treatment, but visible symptoms of water stress were never
apparent in these plants. In contrast, water-stressed ixora
plants (experiment 2, chapter III) exhibited wilting around a
LWP of -1.08 MPa. Therefore, there may have been mechanisms
to maintain turgor under decreasing LWP in plants at the 38
and 43C RZT or plants may have become conditioned to the
high RZTs.
Experiment 2--Growth Room
Banana. Plants held at the 38C RZT under growth room
conditions (Figure 4-4A) had a maximum midday PS rate of 0.81
mg CO2 m-2 s*, which was a 54% increase over the


Figure 4-4. Effects of four root-zone temperatures on the diurnal physiological
responses of 'Grande Naine' banana grown under growth room conditions. A. leaf
photosynthesis B. leaf conductance C. transpiration and D. leaf water
potential. Points are the means of six replicate plants and vertical bars
represent the SE.


A
y-

w 1 jO
28oC
3 3 O C luniiiiiiuiiini
38oC
43oC
, i, - *i
0800 1000 1200 1400 1600 1800
LEAF WATER PO
0
"L > i > '
08 0 0 1000 1200 1400 1600 1800
TIME (hrs)
O


71
corresponding greenhouse rate (Figure 4-2A). The 33C RZT
induced a rate of 0.74 mg CO2 m-2 s-1 at 1000 hr while the
43C RZT treated plants had the lowest PS rate of 0.39 mg CO2
m-2 s_1 at 0800 hr and through the afternoon hours(Figure 4-
4A).
Maximum stomatal opening for the 28 and 33C RZT
occurred earlier in the day than for the higher RZT
treatments (Figure 4-4B). While maximum CS of 0.72 and 0.63
cm s"l occurred in plants at 1000 hr under the 28 and 33C
RZT respectively, peak values for the 43 and 38C RZT were
0.52 at midday and 0.70 cm s-1 at 1400 hr, respectively
(Figure 4-4B). The 43C RZT caused significant reductions in
CS and TR from 0800 hr to 1000 hr and later in the day at
1800 hr. Highest TR rates occurred with the 38C RZT with
values of 94.2 and 92.7 mg H20 m-2 s"1 at 1000 and 1400 hr,
respectively (Figure 4-4C).
Diurnal patterns of LWP were significantly influenced by
RZT treatments (Figure 4-4D). The 28C RZT treated plants had
the highest midday LWP of -0.55 MPa, which was higher than
the LWP at the 33 and 38C RZT treatments. The lowest midday
LWP of -1.26 MPa was induced by the 43C RZT. Overall, LWP
were lower in plants in this experiment than those observed
in the greenhouse environment. This could be attributed to
the higher CS and TR rates that were exhibited by plants in
the growth room experiment compared to the greenhouse. In
both environmental conditions, however, the highest RZT of
43C induced the lowest LWP with higher LWP at lower RZTs.


72
Although the 38 and 43C RZTs appeared to induce water
stress in banana as reflected by low LWP, there was no leaf
folding and/or chlorosis as shown by plants that had been
water stressed to -0.52 and -0.66 MPa by irrigation
treatments under greenhouse and growth room conditions,
respectively (experiments 1,3 Chapter III).
Under growth room conditions, the 33 and 38C RZTs
induced the highest PS rates though at different times of the
day. Under sunlit growth room conditions, 33C air
temperature was reported as being optimum for growth and
partitioning in Cavendish banana but 37C caused leaf injury
(142). Since growth measurements were not recorded in this
study it was impossible to determine whether the increased PS
observed at 38C RZT resulted in increased growth or was
merely a response to the increased root sink demand for
photosynthates. The response of the banana plants grown at
38C RZT in the two environmental conditions would indicate a
greater tolerance to RZTs up to 38C under growth room
conditions. The 43C treatment was supraoptimal under both
environments.
Ixora. Maximum PS, CS and TR at midday followed by
distinct declines were not as evident in ixora plants under
growth room conditions (Figure 4-5) as they were in the
greenhouse study (Figure 4-3). In plants subjected to the 28
and 33C RZT treatments, PS rates of 0.51 and 0.47 mg CO^ m2
sl, respectively, were recorded at 1000 hr and followed by a
gradual decline to 0.21 and 0.12 mg CO2 m-^ s_1,


Figure 4-5. Effects of four root-zone temperatures on the diurnal physiological
responses of Ixora chinenesis L. 'Maui' grown under growth room conditions. A.
leaf photosynthesis FT leaf conductance and C. transpiration. Points are the
means of six replicate plants and vertical bars represent the SE.


PHOTOSYNTHESIS (mg C02 m-2 8-1)
&&


75
respectively, at 1800 hr (Figure 4-5A). A PS rate of 0.57 mg
CO2 m2 S-1 was observed at 0800 hr in the 38C RZT treated
plants and was followed by a significant decline at 1000 hr
with no further decreases for the rest of the day. PS rates
were reduced during the morning hours in plants treated with
the 43C RZT relative to other treatments but increased to
levels comparable to other RZTs at 1400 and 1600 hr. Except
for shifts in time at which maximum daily PS occurred, there
appeared to be little differences in overall PS values
between RZT treatments.
Diurnal CS (Figure 4-5B) and TR (Figure 4-5C) followed
the same general pattern. Maximum values for both parameters
occurred at 1200 and/or 1400 hr in plants treated with either
the 28, 33 or 38C RZT, with subsequent declines for the
rest of the day. Maximum CS recorded for these RZTs was
0.51 cm s"l at 1200 hr which was comparable to the
corresponding midday CS in the greenhouse. However, a CS of
0.46 cm s~l at 1400 hr in the growth room experiment
represented a 302 increase over the CS recorded at this hour
in the greenhouse. A similar pattern was observed for TR.
Maximum TR was 83.3 and 80.1 mg H2O m"2 s-* at 1200 and
1400 hr, respectively. CS and TR were significantly reduced
throughout most of the day in the 43C RZT treated plants
(Figure 4-5B, 4-5C).
Under growth room conditions, therefore, RZTs of 28,
33 and 38C did not produce appreciable differences in
physiological responses in ixora. The relatively low PS, CS


76
and TR early in the day with subsequent increases at later
hours in the 43C treated plants suggested a lag effect of
this RZT on stomata! opening.
Midday CS and TR were comparable to those observed in
the greenhouse. However, higher values for these parameters
were recorded at 1400 hr and later in the growth room as
compared to the distinct post-midday declines in the
greenhouse. There was an apparent effect of the constant high
light intensity during the illumination period in the growth
room that maintained midday stomatal opening to later hours.
However, the eventual decline in gas exchange processes at
1800 hr in the growth room suggested that the stomata may not
have completely lost the diurnal rhythmic pattern evident
under greenhouse conditions.


CHAPTER V
ROOT-ZONE TEMPERATURE AND SOIL MOISTURE EFFECTS ON
BANANA AND IXORA UNDER TWO GROWING CONDITIONS
Introduction
High temperature and water stress often occur
simultaneously in crop situations and alleviating the latter
is often considered a remedy for the former. Although many
drought tolerant plants are also heat tolerant (85),
interactions and sometimes negative correlations occur
between heat and drought stress (102). Research has tended to
concentrate on water and temperature stress effects
separately with little emphasis on defining or separating the
characteristics of these two stress factors.
Suboptimal root-zone temperature (RZT) (7,8,18) and
temperate plant species (5,18,69) have been employed in most
of the reported investigations that involve RZT and soil
moisture interactions. Barlow (7) found that increased shoot
carbohydrate under reduced soil moisture was offset by
increasing RZT in corn seedlings. Kaufmann (69) found that
soil water deficits directly affected leaf water potential
(LWP) and transpiration (TR) in 'Monterey' pine seedlings but
RZT had no consistent effects on TR and CS. Water stress has
77


78
been shown to affect growth (31,116), yield, WUE (10,11,40)
and physiological responses (25,126) in banana (Musa spp.
AAA). Relatively fewer studies have been reported on
temperature stress effects (63,142,143). An air temperature
of 33C was reported as being optimum for growth and dry
weight partitioning in Cavendish banana (142) but the
investigators experienced some difficulty in separating the
stress effects of air temperature and induced vapor pressure
deficits. Investigations in banana involving both stress
factors simultaneously have not been reported. A similar
pattern of research exists for coffee, a major tropical crop
that belongs to the same subfamily as ixora (Ixora chinensis
L. Maui') (6).
This study investigated the effects of four RZTs and two
irrigation volumes on growth, physiology and carbohydrate
status of 'Grande Naine' banana and Maui1 ixora under
greenhouse and growth room conditions.
Materials and Methods
Plant Materials and General Cultural Procedures
Ten- to 12-cm tissue-cultured 'Grande Naine' banana
plants and 12 to 15 cm uniform ixora rooted cuttings were
obtained from commercial nurseries. Plants were hardened in
intermittent mist (6 secs min-1) under 80% light exclusion
for one week and then moved to 40% light exclusion for
another week. Plants were transplanted to 6.5-cm diameter x
25.0-cm tall conical containers (555 cm3) using Metro-Mix 300


79
growth medium (W.R. Grace and Co., Cambridge, MA). Plants
were then moved to the experimental greenhouse or growth room
where they were watered to container capacity daily and
allowed to acclimatize for one week prior to the initiation
of the experiment.
Experiment 1--Greenhouse
Experiment 1 was initiated in August, 1985, in an air-
conditioned greenhouse in which maximum irradiance levels
ranged from 600 to 800 umol m"2 S-1 ppfd, and 25 to 30C day
and 18 to 21C night temperatures were maintained. Relative
humidity was not controlled and varied from 40% to 80%
(Figure 5-1).
Two irrigation volumes and four RZT treatments were
factorially combined in a split plot design experiment.
Plants were watered daily at 2200 hr with 50+5 ml (Wl) and
100+_10 ml (W2) per container ( 555 cm3). These volumes
represented 60% to 70% and 85% to 100% container capacity
(CC), respectively. Water was applied automatically through
drip tubes by a battery-operated controller (Water Watch
Corp. Seattle, WA). The Wl and W2 irrigation treatments were
applied by one and two 6-cm dramm irrigation rings/container,
respectively.
RZTs of 28+_l, 33 +_10, 38+_l and 43+_lC were established
in air-bath boxes as described in Chapter IV. Each
temperature box contained six ixora and six banana plants to
which the irrigation treatments Wl and W2 were randomly


PPFD lumol
Figure 5-1. Photosynthetic photon flux density and relative humidity in the
greenhouse during measurements of physiological parameters.
PERCENT RELRTIUE HliniDITY


81
assigned to three plants per species per treatment.
Temperature boxes were replicated three times. RZT treatments
were applied daily from 1000 to 1600 hr EDT and irrigation
was supplied at 2200 hr so that the imposition of temperature
and irrigation treatments did not overlap. Plants were
fertilized weekly with the appropriate volume of water and
concentrations of 20N-8.8K-16.6P fertilizer (Peters 20-20-20,
W.R. Grace and Co., Cambridge, MA) to apply 250 mg
N/container. Fertilizer was applied with a hand operated
injector (Chem-trol Inc, Kansas City, KS).
Plant height, leaf area, leaf number and stem diameter
for banana, and plant height, plant width, shoot number and
total axillary shoot length for ixora were recorded weekly.
Plant height was considered the distance from the soil
surface to the leaf apex in banana and to the stem tip in
ixora. Banana stem diameter was measured at pot rim level and
leaf area of the third newest leaf was calculated from 0.65
of leaf length multiplied by leaf width (127). Leaf numbers
were derived from the number of whole green functional leaves
and emerging leaves were assigned a value of 0.25, 0.50 or
0.75 depending on the fraction of the leaf lamina fully
exposed.
Ten weeks after the initiation of the experiment, leaf
photosynthesis (PS), leaf conductance (CS) and transpiration
(TR) were recorded on a cloudless day from 1000 to 1600 hr at
2-hr intervals using a portable photosynthesis system (Model
LI-6000, LI-C0R Inc., Lincoln, NE). A 1-liter leaf cuvette


82
was used and the mean of eight 30-sec observations
constituted a measurement. A zero check of the analyzer was
performed between treatments within each replicate.
Simultaneous measurements of LWP were made using a pressure
chamber (PMS Instrument Model 600 Santa Barbara, CA) as
described by Barrett and Nell (9). The third most recently
expanded leaf in banana was selected for gas exchange
processes and LWP measurements. This leaf has been shown to
be the most responsive and the youngest with fully developed
stomata (127). The most recently-matured ixora leaves were
used for PS, CS and TR measurements but 4- to 5- cm shoot
tips were sampled for LWP.
After physiological measurements, leaf and root samples
were immediately collected for carbohydrate analysis. The
third newest leaf in banana and six of the most fully
developed leaves in ixora were sampled. A 5- to 7-g composite
sample of carefully washed roots was used for root
carbohydrate analysis. Plants were then separated into
leaves, stems and roots before drying for at least 48 hr in a
forced-air oven at 70C prior to recording dry weights.
Ethanol-soluble sugars and starch were determined
according to the procedure used by Stamps (133) in which free
sugars were extracted from a 50-mg sample with 80% ethanol.
Leaf and root tissues were first freeze-dried at -40C
(Freezemobi1e II, Virtis Co. Gardner, NY) for at least 72 hr
and stored in a desiccator. Tissue was then ground through a
20-mesh screen (Wiley Mill, Philadelphia, PA) and a 50-mg


83
sample was boiled in 8 ml of 80% ethanol and centrifuged for
1 hr at 10,500 rpm (Model HT Centrifuge, International
Equipment Co., Division of Damon). A 0.5-ml sample of
supernatant was diluted from 10 to 20 times with deionized
water. One half milliliter of 5% phenol and 2.5 ml
concentrated H2SO4 were added to 0.5 ml of each diluted
sample and the mixture was allowed to cool. Absorbance was
then measured at 490 nm using a spectrophotometer. Free sugar
concentrations were determined from plots of absorbance by
standard solutions of D-glucose and deionized water. Starch
content was determined by incubating the centrifuged pellet
at 34C for 12 hr in 5 ml of an enzyme solution containing
sodium phosphate mono- and dibasic, alpha-amylase,
amyloglucosidase and anhydrous calcium chloride. After
incubation, samples were centrifuged for 1 hr and absorbance
determined using the same procedure as above.
The experiment was arranged in a split plot design with
three blocks in which RZT were main plots and irrigation
treatments represented subplots. Each species was treated and
analyzed as a separate experiment. Mean values of
physiological measurements were plotted across time to reveal
diurnal patterns and regression analyses of midday
physiological parameters over RZT were performed for each
irrigation volume treatment. Weekly leaf area data of banana
were used for growth over time plots and final growth and
carbohydrate measurements were used for regression analyses
over RZT and irrigation treatments. Single degree-of-freedom


84
orthogonal comparisons between treatment means were performed
and partitioning of interactions for linear, quadratic and
cubic models for final growth and carbohydrate data were made
using the General Linear Model procedure of the Statistical
Analysis System (134).
Experiment 2--Growth Room
In order to further monitor the physiological responses
of banana and ixora to RZT and irrigation treatments under
more precisely controlled environmental conditions,
experiment 1 was repeated in a walk-in growth room in August
and September, 1986. The growth room was described in
Chapters III and IV. Irrigation and RZT treatments were the
same as for experiment 1 but root heat tube (RHT) containers
as described in Chapter III were used. Similar plant
materials and cultural procedures were used as for experiment
1 except plants were first transplanted for one week into 4-
mi1 7.5 cm diameter x 22.5 cm tall clear plastic bags (1200
cm3) that were inserted into similar sized lengths of PVC
tubes. One week prior to the initiation of treatments, the
plastic bags were transferred to RHTs in the growth room.
Irrigation volumes of 75jj8 ml (Wl) and 150 +_15 ml (W2) per
container were applied daily at 2200 hr as in experiment 1.
These volumes provided 60% to 70% container capacity (CC) and
85% to 100% CC, respectively. Water was applied automatically
by drip irrigation and one and two 1-mm drip tubes per
container constituted the Wl and W2 treatments, respectively.


85
There were 16 RHTs per electronic controller, four of
which were randomly assigned to each RZT of 28 + 0.3, 33+_0.3,
38 + 0.3 and 43+_0.3C. There were six replicate plants per
RZT-irrigation treatment arranged in a randomized complete
block design with three blocks. Species were organized and
treated as separate experiments. RZT treatments were applied
from 1000 hr to 1600 hr daily.
Six weeks later PS, CS, TR and LWP were measured at 2 hr
intervals from 0800 to 1800 hr and growth data and samples
for carbohydrate analysis were taken as described in the
greenhouse experiment. Chlorophyll concentration of leaves
used for physiological measurements was recorded using a Spad
501 chlorophyll light absorbance meter (Minolta Corp. Ramsey,
NJ) and expressed as umol m2.
Means and standard errors were calculated from
physiological measurements of six replicate plants at each
sampling time and are shown with data points. Single degree-
of-freedom orthogonal comparisons between treatment means and
partitioning of interactions for linear, quadratic and cubic
models for growth, physiological and carbohydrate data were
made using the General Linear Model procedure of the
Statistical Analysis System (134).


86
Results and Discussion
Experiment l--Greenhouse
Physiological responses--banana. Under the W1 irrigation
volume, maximum PS at 1200 and 1400 hrs was 0.285 and 0.263
mg CO2 m2 s"1, respectively (Figure 5-2A). Diurnal patterns
for CS and TR appeared to parallel each other under both
irrigation levels with maximum levels for both parameters
occurring at 1200 and 1400 hr (Figure 5-2, 5-3). LWPs were
generally higher in plants at the 28 and 33C RZTs under the
W1 irrigation volume except at midday when LWP averaged
-0.71 MPa over all RZTs (Figure 5-2D).
At the higher irrigation level, W2, a maximum PS of
0.330 mg CO2 m-2 s was measured for the 33C RZT treated
plants at midday (Figure 5-3A) and another similar peak of
0.301 mg CO2 m-2 s* occurred in the 38C RZT treated plants
at 1400 hr. PS was significantly reduced at midday and
thereafter by the 43C RZT. LWPs were generally higher in the
33 and 38C RZT treated plants compared to those observed at
28 and 43C between 1000 and 1400 hr (Figure 5-3D).
Regression analyses of midday PS over RZT and irrigation
treatments revealed significant effects of RZT but not
irrigation treatments (Figure 5-4A). PS was highest at the
28 and 33C RZT treatments with progressive reductions at
the 38 and 43C RZTs. While highest midday PS was noted for
the 28 and 33C RZTs, diurnal patterns showed a lag in
response for the 38C RZT with peak PS occurring after midday
(Figure 5-2A, 5-3A). Since there were also corresponding


Figure 5-2. Effects of four root-zone temperatures and a 50+5 ml daily
volume per 555 cm3 container on the diurnal physiological responses of
Naine* banana grown under greenhouse conditions. A. leaf photosynthesis
conductance C. transpiration and D. leaf water potential. Points are
of six replicate plants and vertical bars represent the SE.
irrigation
'Grande
B. leaf
the means


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ROOT-ZONE TEMPERATURE AND SOIL MOISTU
ON GROWTH AND PHYSIOLOGY
OF CONTAINER-GROWN 'GRANDE NAINE'
AND IXORA CHINENSIS L. 'MAU
RE EFFECTS
BANANA
r
By
CHRISTOPHER RAMCHARAN
A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA
IN PARTIAL FULFILLMENT OF THE REQUIREMENTS
FOR THE DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1987

ACKNOWLEDGEMENTS
I want to especially thank my major advisor, Dr.
Dewayne. L. Ingram. He impressed upon me the art of
scientific writing, an enthusiasm for research and the will
to persevere. Also, I wish to thank the other members of my
committee, Drs. Terril Nell, James Barrett, Jerry Bennett and
William Wiltbank, for their encouragement and critical
evaluation of my work.
I am deeply grateful to the United States Department of
Agriculture whose financial support and grant made this
research possible and to the Department of Ornamental
Horticulture for support staff, equipment and services. The
contributions of Ms. Claudia Larsen for laboratory and
technical assistance and T.D. Townsend of the greenhouse
staff are highly appreciated.
I would like to thank Oglesby nursery for the generous
donations of banana plants and Mr. George Behren of Behren's
nursery for the ixora plants used in this study.
Last, but certainly not least, sincere thanks go to my
family. The support of my wife Senovia throughout the course
of these studies was invaluable. Finally, I wish to
acknowledge my children: they gave me the purpose for which
it was done.

TABLE OF CONTENTS
Page
ACKNOWLEDGEMENTS ii
LIST OF TABLES v
LIST OF FIGURES vii
ABSTRACT xi
CHAPTERS
I GENERAL INTRODUCTION 1
II LITERATURE REVIEW 4
Introduction 4
Water Stress Physiology 5
Physiological Effects of Root-Zone
Temperature 12
Root-Zone Temperature and Soil Moisture
Interactions 18
IIIEFFECTS OF IRRIGATION VOLUME ON BANANA
AND IXORA UNDER TWO GROWING CONDITIONS 22
Introduction 22
Materials and Methods 23
Plant Materials and General Cultural
Procedures 23
Greenhouse 23
Experiment 1 23
Experiment 2 26
Growth Room 26
Experiment 3 26
Results and Discussion 27
Greenhouse 27
Experiment l--banana 27
Experiment l--ixora 34
Experiment 2--banana 40
Experiment 2--ixora 42
Growth Room 42
Experiment 3--banana 42
Experiment 3--ixora 50
i i i

IVROOT-ZONE TREMPERATURE EFFECTS ON BANANA AND
IXORA UNDER TWO GROWING CONDITIONS 56
Introduction 56
Materials and Methods 57
Plant Materials and General Cultural
Procedures 57
Experiment l--Greenhouse 57
Experiment 2--Growth Room 60
Results and Discussion 61
Experiment l--Greenhouse 61
Banana 61
Ixora 65
Experiment 2--Growth Room 68
Banana 68
Ixora 72
VROOT-ZONE TEMPERATURE AND SOIL MOISTURE
EFFECTS ON BANANA AND IXORA UNDER TWO
GROWING CONDITIONS 77
Introduction 77
Materials and Methods 78
Plant Materials and General Cultural
Procedures 78
Experiment l--Greenhouse 79
Experiment 2--Growth Room 84
Results and Discussion 86
Experiment l--Greenhouse 86
Physiological responses--banana .... 86
Growth responses--banana 93
Carbohydrate analysis--banana 100
Physiological responses--ixora 104
Growth responses--ixora 109
Carbohydrate analysis--ixora 118
Experiment 2--Growth Room 120
Physiological responses--banana .... 120
Growth responses--banana 128
Carbohydrate ana 1ysis--banana 132
Physiological responses--ixora 137
Growth responses--!* xora 145
Carbohydrate analysis--ixora 150
VISUMMARY AND IMPLICATIONS 153
LITERATURE CITED 160
BIOGRAPHICAL SKETCH 172

LIST OF TABLES
Page
5-1. Growth components of 'Grande Naine' banana
measured after 10 weeks at four root-zone
temperatures and two irrigation volumes under
greenhouse conditions 94
5-2. Dry weight components of 'Grande Naine' banana
measured after 10 weeks at four root-zone
temperatures and two irrigation volumes under
greenhouse conditions 98
5-3. Shoot and root carbohydrate distribution of
'Grande Naine' banana at four root-zone
temperatures and two irrigation volumes under
greenhouse conditions 101
5-4. Growth components of Ixora chinensis L.'Maui'
measured after 10 weeks at four root-zone
temperatures and two irrigation volumes under
greenhouse conditions 112
5-5. Shoot and root carbohydrate distribution of
Ixora chinensis L. 'Maui' measured after 10 weeks
at four root-zone and two irrigation volumes
under greenhouse conditions 119
5-6. Growth components of 'Grande Naine' banana
measured after six weeks at four root-zone
temperatures and two irrigation volumes under
growth room conditions 129
5-7. Dry weight components of 'Grande Naine' banana
measured after six weeks at four root-zone
temperatures and two irrigation volumes under
growth room conditions 130
5-8. Shoot and root carbohydrate distribution of
'Grande Naine' banana at four root-zone
temperatures and two irrigation volumes under
growth room conditions 134
v

5-9. Growth components of Ixora chinensis L.'Maui*
measured after six weeks at tour root-zone and
two irrigation volumes under growth room
conditions 146
5-10. Dry weight components of Ixora chinensis L.'Maui'
measured after six weeks at four root-zone
temperatures and two irrigation volumes under
growth room conditions 147
5-11. Shoot and root carbohydrate distribution of
Ixora chinensis L. 'Maui' measured after six weeks
at four root-zone temperatures and two irrigation
volumes under growth room conditions 151

LIST OF FIGURES
Page
3-1. Photosynthetic photon flux density and relative
humidity in the greenhouse during measurements
of physiological parameters 24
3-2. Effects of three irrigation volumes on the
diurnal physiological responses of 'Grande
Naine' banana under greenhouse conditions 29
3-3. Effects of three irrigation volumes on the
diurnal water use efficiency of 'Grande Naine'
banana under greenhouse conditions 33
3-4. Effects of three irrigation volumes on the
diurnal physiological responses of Ixora
chinensis L. 'Maui' under greenhouse
conditions 36
3-5. Effects of three irrigation volumes on the
diurnal water use efficiency of Ixora
chinensis L. 'Maui' under greenhouse
conditions 39
3-6. Effects of a 14 day drying cycle on leaf water
potential and leaf conductance of 'Grande Naine'
banana under greenhouse conditions 41
3-7. Effects of a 14 day drying cycle on leaf water
potential and leaf conductance of Ixora
chinensis L. 'Maui'under greenhouse
conditions 43
3-8. Effects of three irrigation volumes on the
diurnal physiological responses of 'Grande Naine'
banana under growth room conditions 45
3-9. Effects of three irrigation volumes on the
diurnal water use efficiency of 'Grande Naine'
banana under growth room conditions 49
vi i
I

3-10. Effects of three irrigation volumes on the
diurnal physiological responses of Ixora
chinensis L. 'Maui' under greenhouse
conditions 52
3-11. Effects of three irrigation volumes on the
diurnal water use efficiency of Ixora
chinensis L. 'Maui ' under growth room
conditions 55
4-1. Photosynthetic photon flux density and relative
humidity in the greenhouse during measurements of
physiological parameters 58
4-2. Effects of four root-zone temperatures on the
diurnal physiological responses of 'Grande
Naine ' banana grown under greenhouse
conditions 63
4-3. Effects of four root-zone temperatures on the
diurnal physiological responses of Ixora
chinensis L. 'Maui' grown under greenhouse
conditions 67
4-4. Effects of four root-zone temperatures on the
diurnal physiological responses of 'Grande
Naine' banana grown under growth room
conditions 70
4-5. Effects of four root-zone temperatures on the
diurnal physiological responses of Ixora
chinensis L. 'Maui' grown under growth room
conditions 74
5-1. Photosynthetic photon flux density and relative
humidity in the greenhouse during measurements of
physiological parameters 80
5-2. Effects of four root-zone temperatures and a 50+_5
ml daily irrigation volume per 555 cm3 container
on the diurnal physiological responses of 'Grande
Naine' banana grown under greenhouse
conditions 88
5-3. Effects of four root-zone temperatures and a 100+_10
ml daily irrigation volume per 555 cm3 container
on the diurnal physiological responses of 'Grande
Naine' banana grown under greenhouse
conditions 90
5-4. Regressions of midday physiological responses of
container-grown 'Grande Naine' banana over
four root-zone temperatures and two irrigation
volumes under greenhouse conditions 92
v i i i

5-5. Effects of four root-zone temperatures across
two irrigation volumes on leaf area of the third
newest leaf in container-grown 'Grande Naine'
banana measured over 10 weeks under greenhouse
conditions 96
5-6. Effects of four root-zone temperatures and two
irrigation volumes on shoot/root ratio and root
dry weight of container-grown 'Grande Naine'
banana grown under greenhouse conditions 99
5-7. Effects of four root-zone temperatures and two
irrigation volumes on root sugar/starch content
of container-grown 'Grande Naine' banana
under greenhouse conditions 103
5-8. Effects of four root-zone temperatures and a 50+_5
ml daily irrigation volume per 555 cm3 container
on the diurnal physiological responses of Ixora
chinensis L. 'Maui' grown under greenhouse
conditions 106
5-9. Effects of four root-zone temperatures and a 100+_10
ml daily irrigation volume per 555 cm3 container
on the diurnal physiological responses of Ixora
chinensis L. 'Maui' grown under greenhouse
conditions 108
5-10. Regressions of midday physiological responses of
container-grown Ixora chinensis L. 'Maui' over
four root-zone temperature and two irrigation
volumes under greenhouse conditions Ill
5-11. Main effects of four root-zone temperatures
across two irrigation volumes on growth of Ixora
chinensis L. 'Maui' grown under greenhouse
conditions 115
5-12. Effects of four root-zone temperatures and two
irrigation volumes on dry weight components of
container-grown Ixora chinensis L. 'Maui'
grown under greenhouse conditions 117
5-13. Effects of four root-zone temperatures and a 75+_8
ml daily irrigation volume per 1200 cm3 container
on the diurnal physiological responses of
container-grown 'Grande Naine' banana grown under
growth room conditions 122
5-14. Effects of four root-zone temperatures and a 150+_15
ml daily irrigation volume per 1200 cm3 container
on the diurnal physiological responses of container-
grown 'Grande Naine' banana grown under
growth room conditions 124
i x

5-15. Regressions of midday physiological responses
of container-grown 'Grande Naine' banana over
four root-zone temperatures and two irrigation
volumes under greenhouse conditions 127
5-16. Effects of four root-zone temperatures across
two irrigation volumes on chlorophyll
concentration of container-grown 'Grande Naine'
banana under growth room conditions 133
5-17. Interactive effects of four root-zone temperatures
and two irrigation volumes on carbohydrate content
of container-grown 'Grande Naine' banana
under growth room conditions 137
5-18. Effects of four root-zone temperatures and a 75+8
ml daily irrigation volume per 1200 cm3 container
on the diurnal physiological responses of
Ixora chinensis L. 'Maui' grown under growth
room conditions 140
5-19. Effects of four root-zone temperatures and a 150+15
ml daily irrigation volume per 1200 cm3 container
on the diurnal physiological responses of
Ixora chinensis L. 'Maui' grown under growth
room conditions 142
5-20. Regressions of midday physiological responses
of container-grown Ixora chinensis L. 'Maui'
over four root-zone temperatures and two
irrigation volumes under growth room
conditions 144
5-21. Effects of four root-zone temperatures and two
irrigation volumes on growth of container-grown
Ixora chinensis L. 'Maui' under growth room
condi tions 149
x

Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy
ROOT-ZONE TEMPERATURE AND SOIL MOISTURE EFFECTS ON GROWTH
AND PHYSIOLOGY OF CONTAINER-GROWN 'GRANDE NAINE'
BANANA AND IXORA CHINENSIS L. 'MAUI'
By
Christopher Ramcharan
May 1987
Chairman: Dewayne L. Ingram
Major Department: Horticultural Science
Studies to investigate the effects of root-zone
temperature (RZT) and irrigation volume (IRV) on growth and
physiological responses of 'Grande Naine' banana (Musa spp.
AAA) plantlets and Ixora chinensis L. 'Maui' were conducted
under greenhouse (GH) and growth room (GR) environments.
Leaf photosynthesis (PS), conductance (CS) and
transpiration (TR) in the GH were reduced 65%, 57% and 49%,
respectively, in banana under an IRV of 20+_4 ml daily per 150
cm3 container as compared with the highest IRV of 40+8 ml
daily per container. Measured parameters were severely
reduced at an IRV of 10+2 ml per container. Physiological
responses of ixora were also reduced by decreased IRV. Leaf
wilting occurred at a leaf water potential (LWP) of -1.1 MPa
and leaves abscised below a LWP of -2.1 MPa.
xi

In an RZT study, maximum midday PS in banana occurred at
the 33°C RZT in a GH and a maximum midday PS of 0.74 mg CO2
m"2 s'1 was observed in 33° and 38°C RZT-treated plants in a
GR. In ixora, a 33°C RZT induced a maximum midday PS in the
GH but there were no apparent differences in PS between 28°,
33° and 38°C RZTs in the GR study.
Maximum PS in banana occurred at 38° and 33°C RZTs under
a higher IRV of 100+_10 ml daily in the GH. In the GR, maximum
PS occurred in 33°C RZT-treated plants and the higher IRV but
at 38°C RZT under a moderate IRV (75+8 ml daily). Increasing
RZT reduced leaf area under both IRVs. Carbohydrate (CHO)
partitioning between shoot and root was correlated with RZT-
induced changes in LWP.
Gas exchange processes in ixora were reduced by RZTs
above 33°C RZT regardless of IRV. RZT-induced shoot growth
was not associated with increased PS or TR but a hormonal
role was theorized. CHO partitioning was likely related to
the induced shoot growth and was influenced by environmental
condi ti on s.
These findings could have significant applications to
banana production schemes using tissue-cultured plants. RZTs
above 33°C could reduce the landscape and floricultural value
of container-grown ixora.
xi i

CHAPTER I
GENERAL INTRODUCTION
Temperature and water are perhaps two of the most
important environmental factors affecting growth and
physiology of plants. Although soil moisture can be modified
by irrigation, altering temperature is more difficult.
Extreme temperatures produce deleterious but often subtle
effects. Soil temperature and water are interrelated since
both directly affect plant roots. Maintaining optimum water
relations and temperature may be difficult, especially with
container-grown species.
Hot and dry conditions often occur simultaneously in
crop situations and alleviating the latter is often
considered a remedy for the former. Although many drought
tolerant plants are heat tolerant (85), the interactions
between temperature and water stresses have not been well
defined (102). Suboptimal root-zone temperatures (RZT) stress
plants through reduced respiration, increased water viscosity
(82) and reduced absorption caused by decreased cell membrane
permeability (74). In this context, increased RZTs, within
limits, are believed to have the reverse effect and so
increase water (5,30) and possibly nutrient absorption
(39,44).
1

2
Within limits, increased soil temperatures have
generally increased shoot growth curvi1inear 1y (30).
Restricted shoot or leaf elongation is usually the first
symptom of mild water stress (57). Reduced soil water might
offset increased shoot growth induced by increasing soil
temperature to within the optimum range. A similar induction
of increased nutrient absorption by increasing soil
temperature may occur thus permitting less than normal
nutrient application. Such a recommendation was made for
peppers (Capsicum annuum L.) grown at a supraoptimal RZT of
360c (44).
Water stress, by its immediate effect on shoot growth
but delayed action on photosynthesis (13), has been shown to
increase shoot carbohydrate concentrations (8,119). Increased
root temperature has also been shown to reduce shoot
carbohydrates (7), again suggesting the possibility of
interactive effects between the two stress factors. Indeed
some pasture grasses were of better quality because of higher
carbohydrate levels under high temperature and dry conditions
as opposed to being irrigated (18).
Interactive effects of soil water and RZT on
translocation and carbohydrate partitioning patterns have
been reported (7,8). Increased RZTs have altered
translocation rates due to increased sink demands of roots
(46). Water stress could reduce translocation by its effect
on reduced turgor pressure and the consequent effect on
decreased mass flow of assimilates.

Therefore, the overall objective of this study was to
identify the effects of supraoptimal RZT and water deficits
on two container-grown plants. The specific objectives were
to
1) Determine growth and physiological effects on 'Grande
Naine' banana (Musa spp. AAA) and ixora (Ixora chinensis L.
'Maui') and to elucidate whether the two stress factors act
independently or interactively.
2) Identify any compensatory type interactions that may
be exploited in container production of plants.
3) Determine if drought stress can be induced by high
RZT, identify critical RZTs and observe whether watering
offsets temperature-induced water stresses.
Banana and ixora were selected because they represent a
contrast in morphology, phenotype and horticultural usage.
The use of tissue-cultured banana plants in this study
represents a unique application of micropropagation as a tool
in plant physiology. The choice of banana was also in
anticipation of the stress problems that may arise in the
banana production schemes that convert to the use of tissue-
cultured plantlets.

CHAPTER II
LITERATURE REVIEW
Introduction
Temperature and water stress affect plant growth and
physiological processes. The literature is abundant with data
describing their independent effects but there have been few
attempts to separate the effects of these two stress factors
when applied in combination. Any attempt to determine
interactions between soil water and temperature on plants
must first review or determine their independent effects.
While it is generally accepted that water stress reduces
expansive growth and induces a progressive reduction in gas
exchange processes, the effects of supraoptimal RZTs have not
been clearly identified. Soil water deficits usually reduce
plant water status and supraoptimal RZTs may affect root
absorption and conductance and indirectly stomatal function.
However, plant water stress can occur even with adequate soil
water when ambient conditions favor excessive transpiration
or water absorption is inhibited by factors including high
RZT.
Since supraoptimal root temperature directly affects
root respiration and metabolism, its effect on partitioning
4

5
patterns within the plant seems likely more direct than that
of water stress.
Water stress physiology
Water stress basically results from an imbalance
between soil water absorption and that required by plants to
meet transpirationa1 needs. Kramer (74) stated that plant
water stress is caused by either excessive water loss,
inadequate absorption or a combination of the two. In
general, an increase in water stress produces a two phase
response in photosynthesis, with a threshold water potential
above which there is little or no change in photosynthesis
and below which the rate decreases rapidly (114). A water
potential of -0.4 MPa reduced the photosynthetic capacity of
loblolly pine with assimilation being negligible at -1.1 MPa
(17). This effect was caused by stomatal closure which
reduced transpiration as well as photosynthesis. Water stress
may affect photosynthesis via three major mechanisms (131):
1) by stomatal closure, 2) by cellular dehydration which
reduces enzyme activity, and 3) by decreasing cell membrane
permeability to carbon dioxide and bicarbonate.
Although stomatal closure has been cited as the primary
method by which water stress affects photosynthesis
(13,78,132), there are several reports of non-stomatal
reductions in photosynthesis (14,23,49,114). Many species
that exert some degree of stomatal limitation on
photosynthesis at low water potentials characteristically
have high mesophyll resistance (50,141). When the components

6
of leaf resistance to CO2 were measured for several species,
mesophyll resistance was found to be lower than stomatal
resistance but about ten times that of boundary layer
resistance (139). Soil water deficit also affects the diurnal
pattern of stomatal aperture (33) with progressi ve 1y earlier
closing under prolonged drought conditions (93).
Stomatal closure affected photosynthesis mainly by its
effect on the carbon dioxide supply (13,17). However, some
researchers agree that photosynthesis is evident even when
stomata are closed (93). Because of their diffusive nature,
photosynthesis and transpiration are both affected by
stomatal closure, although transpiration may be affected more
than photosynthesis (48,120). Photosynthesis was reduced 25%
while transpiration decreased 50% in apple (Mal us pumi 1 a
Mill.) under drought conditions (51). Zelitch (149) indicated
that partial stomatal closure reduced transpiration
relatively more than photosynthesis. Although the diffusive
resistance of CO2 was larger than that of water, CO2 also
encountered a vastly greater mesophyll resistance than water.
Consequently, the stomata represented a smaller portion of
the total resistance to CO2 than to water vapor.
Moorby et al. (95) found no effect of drought on the
activity of ribulose 1,5-diphosphate (RUDP) carboxylase or
carbonate dehydratase, two enzymes involved in CO2 fixation.
Ackerson et al. (1) however, reported that water stress
reduced the activity of photosynthetic enzymes of potato
(Solanum tuberosum L). Lawlor (84) indicated that ATP

7
synthesis through photophosphorylation was inhibited by water
stress while electron transport and reductant supply were
relatively insensitive. Boyer (14) demonstrated that
photosynthesis of water stressed sunflower (Helian thus annuus
L.) plants under growth chamber conditions was affected more
by photochemical activity than by CO2 diffusion or enzymatic
activity. Davies (33) showed that stomatal sensitivity to
decreasing leaf water potential in cotton (Gossypi urn hirsutum
L.) and soybean (Glycine max. L.) was greater for plants
grown in a growth chamber than in a greenhouse.
When soil moisture is adequate, diurnal fluctuations in
xylem pressure potential closely followed the daily trend of
atmospheric evaporative demand, whether expressed as
radiation, vapor pressure or humidity (22,71,121). As soil
moisture becomes limiting, early morning leaf water
potentials (LWP) begin at a lower value, decrease sharply and
then remain constant over several hours, reflecting the
influence of increased stomatal resistance (29,108).
Threshold relationships between diffusive resistance and leaf
water potential appear to be more common than linear or
curvilinear relationships (52,67,68). Brix (17) showed that
net assimilation rate (NAR) was closely related to LWP; NAR
began to decrease at -1.0 MPa and completely ceased at -3.5
MPa. Other reports (53,79,113) indicated that parallel
decreases in NAR and LWP were strongly linked to changes in
diffusive resistance. However, NAR changes in pear (Pyrus
communis L.) (77), Pacific silver fir (Abies alba Mill.) and

8
noble fir (Abies nobi 1 is Lindl.) (55) did not always coincide
with changes in diffusive resistance indicating the influence
of mesophyll and or carboxylation resistance. It was
concluded that species vary considerably in their
physiological reactions to water deficits.
The majority of studies on water stress in banana have
dealt primarily with the effects on fruit bunch and yield
characteristics, but with few details reported on
morphological, phenological and physiological responses.
However, all reports agree that water stress causes a general
reduction in growth (19,111,136). Daniells (31) in Australia
indicated a dramatic reduction in growth, delayed bunch
emergence, inferior fruits and longer maturity time for
'Williams' banana under decreasing soil water levels.
Robinson and Albertson (116) reported that optimum growth and
yields were obtained by maintaining soil moisture levels at
16 to 34% of total available soil moisture (ASM).
Bhattacharyya and Rao (10) also reported optimum bunch and
yield character!' sti cs with 20 to 40% ASM depletion levels. An
unimpeded supply of water was essential for development of an
efficient root system, rapid rate of leaf emergence,
continued production and partitioning of assimilates, and
synthesis of growth hormones such as cytokinins. In Honduras,
Ghavami (40) found an irrigation rate of 44 mm/week or a soil
moisture tension of 30 to 40 centibars resulted in the
highest yields of 'Valery' banana.

9
Perhaps the most detailed description of the effects of
soil water levels on banana physiology was recorded by
Shmueli (126) in his paper on the physiological activity of
banana in relation to soil moisture in Israel. Depletion of
soil moisture below a third of the available water capacity
was accompanied by marked yellowing of leaves and significant
reduction in stem diameter. Diurnal changes in stomatal
activity were clearly related to ASM. When soil moisture
decreased below two-thirds of total ASM, there was a
reduction in stomatal conductance which was confined mainly
to the morning hours. The impairment of plant water balance
became acute when soil moisture decreased to about one-third
of the total available. Although diurnal patterns of stomatal
conductance and transpiration generally paralleled each
other, the rate of transpiration was considerably more
influenced by environmental conditions. Soil moisture levels
below two-thirds of field capacity resulted in reduced leaf
osmotic potential. Overall, banana plants responded to
diminishing soil moisture by stomatal closure, reduced
assimilation and accelerated yellowing of leaves indicative
of starvation effects.
Current efforts to increase agricultural production,
coupled with conservation of limited high quality water, have
focused attention on the possibility of increasing water use
efficiency (WUE) of various crops (16,88,97). Fischer and
Turner (38) observed that under field conditions, WUE usually
remained constant or increased slightly with increasing soil

10
water limitations for C3 species. Reports on the effects of
water stress on WUE are conflicting. Some investigations
pointed to only small increases in WUE with water stress
(16,38,97). Reduction of water supply to wheat
(Triticum aestivum L.) such that transpiration was reduced by
70% of the controls was associated with a 20% increase in WUE
(41). On the other hand, WUE in maize (Zea mays L.) was lower
on days when there was increasing soil water stress (128).
Limited soil water resulted in yield losses but enhanced WUE
in grapefruit (Citrus paradisi L.) (16) and Shamouti orange
(Citrus sinensis Osb.) (97). Manning et al. (86) found a
positive correlation of WUE with soil moisture regime, plant
height, leaf area and seed yield in pea [Pisum sativum (L.)
Mer. ].
Adequate irrigation was critical for vegetative growth
in coffee (Coffea arabica L.) (4,72) and water stress reduced
extension growth, node number and leaf area (138). Ten and 25
ml water/day/coffee seedling in 15 cm pots resulted in
drastic growth reduction, wilting and necrosis of the
terminal shoots compared to a 200 ml treatment which caused
water logging and induced leaf abscission (72).
Water stress has been shown to cause specific changes in
plant carbohydrate status. Decreased starch concentration
simultaneous with increased sugar content during drought
stress have been reported for sugar maple (Acer saccharum
Marsh.) (106) and black oak (Quercus velutina Lam.) (107). In
sugar maple total carbohydrates were not affected, indicating

11
a change in carbohydrate partitioning. Similar changes in
carbohydrates occurred in the inner bark of loblolly pine
(Pinus taeda L.) (56) and in cotton shoots (36). Water stress
induced increases in sugar concentration may serve several
purposes, of which two are of major significance. Increased
sugars increase the readily available substrate for
respiration (17). A second effect of increased free sugars
during water stress is to lower the osmotic potential of the
cell solution and hence maintain cell turgor. Concentration
and form of carbohydrates, organic acids or inorganic ions
are modified to promote osmotic regulation (74). A more
negative leaf solute potential with water stress was observed
in seedlings of English oak (Quercus robur L.) and silver
birch (Betula pendula Rodt.) but the degree of osmotic
regulation varied between species (105).
Munns and Pearson (98) found that reduced photosynthesis
as a result of water stress was not caused by accumulation of
photosynthates in potato leaves. Low leaf water potential
resulted in decreased translocation of carbohydrate, which
was proportional to the decline in net photosynthesis,
irrespective of whether tubers were present. A lower
proportion of labelled 1^C02 was found in tubers of stressed
potato plants compared to non-stressed plants (125).
One of the earliest plant responses to water stress is
reduced elongation of leaves, stems and roots (57).
Boyer (13) found that leaf enlargement of corn and soybean

12
was inhibited earlier and more severely than photosynthesis
and respiration by decreasing water potentials.
Physiological Effects of Root-Zone Temperature
According to Levitt (85) the two major forms of stress
injury are direct and indirect injury. Direct heat injury
results from a short exposure to an extreme temperature and
is detectable immediately at the cellular level. Indirect
injury results from prolonged exposure to temperatures below
those causing direct injury.
Supraoptimal temperature stress may be defined as the
retardation or cessation of metabolic functions in response
to high temperatures. Alexandrov (3) reported that the first
symptom of high temperature was the cessation of protoplasmic
streaming. This was followed by a reduction in photosynthesis
with subsequent damage to the chioroplasts. Finally,
semipermeabi1ity of cell membranes was disrupted, so that
cellular compartmenta1ization was lost with mixing of
cellular contents.
Starvation, a common form of indirect heat injury in
plants, occurs because of a higher optimum temperature for
respiration than photosynthesis (85). The temperature at
which photosynthesis and respiration rates are equal is the
temperature compensation point. When plant temperatures
exceed this critical point, carbohydrate reserves become
depleted and starvation ensues. Other forms of indirect
metabolic injury (85) are toxicities resulting from

13
disturbance of specific metabolic processes and biochemical
lesions resulting from the accumulation of metabolites
necessary for plant growth.
Ingram and Buchanan (61,62) demonstrated that electrol¬
yte leakage through root cell membranes in response to
increasing supraoptimal temperatures was sigmoidal and the
midpoint of this response was defined as the critical
temperature. Critical temperature was species and cultivar
specific and varied from 48° to 53°C for a 20 minute exposure
time (61,62). Direct injury at high temperature was also
shown to be a function of exposure time (60).
Proteins are denatured by high temperature (2) and there
is evidence that high temperature disrupts membrane
structure. Inhibition of photosynthesis at high temperature
was correlated with the disruption of membrane associated-
processes including photosystem II (108),
photophosphorylation (123) and the change of chlorophyll
organization (122). The stability of membranes is mainly due
to the degree of fatty acid saturation which is linked to
membrane function and organelle association (145).
Environmental temperatures have been demonstrated to alter
fatty acid saturation ratios and membrane fluidity as
adaptive responses to high and low temperatures
(24,27,28,45,87).
There are several reports of the effects of growth
medium temperature on water uptake by roots (70,89,104,115).
Water conductivity through roots appears to be a temperature-

14
dependent process (117). Low temperature reduced
transpiration and stomatal conductance in chi 11-susceptible
species by reducing conductance through root cell membranes
(75,104,147). Such results are most likely due to increased
resistance to water at the endodermis layer (89).
Heating the root-zone to temperatures within the optimum
range generally improved plant growth (12,42,83) and heating
the root medium to 20°C improved foliage plant production at
low greenhouse air temperatures (12). Root temperatures above
or below optimum may alter hydraulic conductivity and thus
indirectly affect stomatal conductance, photosynthesis and
subsequent carbohydrate translocation (66). Optimal root-zone
temperature in tomato (Lycopers icon esculentum Mill.) ranged
from 30° to 36°C for nutrient translocation and water
absorption (42,58).
Using plants with heat-killed roots, Kramer (73) showed
that plant shoots remained alive and did not wilt for several
days even after root death. Transpirati on decreased after
root death because of leaf injury and gum deposits from dead
cells. Citrus hydraulic root conductance increased linearly
with increasing root temperature (137) and Carrizo citrange
exhibited a log-linear decrease in conductivity over a
temperature range from 40° to 10°C (146).
Cooper (30) published an extensive literature review in
1973 on RZT effects and reported important differences due
not only to species and cultivars but also to growing
conditions. High leaf photosynthetic rates in soybean grown
r

15
at increasing RZT were related to low stomatal resistance,
high transpiration rate and high phosphoenol pyruvate and
RUDP carboxylase content (35). Cucumber (Cucumis sativus L.)
leaves were larger, contained more chlorophyll and produced
greater amounts of ATP and NADPH when plants were grown at
25°C RZT compared to 15° or 35°C (26). Canopy photosynthesis
of pepper plants was closely related to shoot dry weight and
crop yield (44), although maximum growth and yields were
obtained at 30°C RZT and maximum photosynthesis rates were
recorded at 36°C RZT. The disparity between optimal RZTs was
explained by increased rates of respiration at the higher
RZT. High rates of photosynthesis at high RZT in tomato were
due not only to increased plant size but also to a
modification of the physiological and morphological propert¬
ies of the photosynthetic apparatus (43). Chlorophyll content
(26), enzyme activity (35) and leaf hormonal content (91)
have been influenced by increasing RZT.
Although many physiological processes are influenced by
high RZTs (30), there is relatively little information on the
effects of RZT on endogenous hormone levels (76). High RZTs
have been reported to increase gibberellin and decrease
cytokinin translocated in the root exudates (47,64,129).
There is also evidence that high RZT may affect the
distribution of gibberellin in the shoot (76). Alternatively,
high RZT may have increased the activity in the roots of
gibberellins previously synthesized in the shoot as well as
influenced synthesis in the shoot (110).

16
In a study on flowering in coffee (92), ambient
temperatures above 33°C induced formation of orthotropic
shoots and prevented flower initiation. Root-zone heating
delayed flowering and induced excessive vegetative growth in
peppers (44). Such an antagonism between vegetative and
reproductive growth prompted the authors to recommend a
reduction in night temperature or nitrogen fertilization when
pepper plants are grown at high RZTs.
Reduced biosynthesis of cytokinins induced by supraopti-
mal RZT was thought to reduce photosynthetic rates in bean
(Phaseolus vulgaris L.) (21) and grape (Vitis vi nifera L.)
(129). Increasing RZTs from 18° to 30°C in rice (Oryza sativa
L.) increased the translocation and rate of
photosynthetically assimilated l^C into the roots but
decreased root growth apparently by retarding protein and
cell wall synthesis (96).
Temperature has long been recognized as the main
environmental determinant of plant growth and phenology (81).
Increased soil temperatures have generally increased shoot
growth curvi1inearly and reduced root growth (30,112). All
growth variables of Pittosporurn tobira Thunb. were
substantially lowered by a 40°C RZT for 6 hrs/day compared to
27°C (65), and Ingram (59) noted marked growth inhibition of
woody plants stressed by 35° to 40°C RZT for 6 hrs/day.
Gosselin and Trudel (44) reported maximum leaf area in
pepper at a RZT of 30°C with a decline at higher
temperatures. Watts (144) demonstrated reduced leaf expansion

17
in maize with increasing RZT but could not explain this as a
result of increased plant water deficits. RZTs above 37°C
restricted leaf growth and rate of emergence in pearl millet
(Pennisetum typhoides S&H) (103). Soil temperatures of 37°C
and above, which are common in the tropics, reduced both root
and shoot growth of maize and cowpea (Vigna unguiculata L.)
(90). Well-watered potted eggplants (Sol an urn melongena L.)
exhibited decreased plant growth with increasing RZT from 25°
to 40°C (118). Shoot to root ratio, however, was constant
except at 40°C, where root rot occurred.
Philpotts (109) reported a linear decrease in cowpea
nodulation and total plant dry weight with increasing RZT
from 31° to 40°C. RZTs above 32°C significantly reduced
vegetative growth of two cultivars of cowpea through their
effects on shoot, peduncle and root dry weight and, to a
lesser extent, leaf number (94).
Using a system of circulating water coils, Franco
(39) applied RZTs from 13° to 48°C and observed the effects
on coffee growth, transpiration and mineral absorption.
Maximum transpiration occurred at 33°C with a significant
decline at 43°C. Similarly, RZTs above 33°C reduced
absorption of several nutrients and induced leaf chlorosis.
Plants growing at 48°C died and both shoot and root growth
were depressed above 33°C. In another experiment using young
coffee seedlings grown at 33°C RZT, he reported the
occurrence of small tumors at the base of the stem from which
new orthotropic or non-flowering shoots grew.

18
In an extensive investigation using 'William' banana
in sunlit growth chambers, Turner and Lahav (142) reported
heat injury at an air temperature of 37°C. Total plant weight
was greatest at 28°C while leaf area was maximum at 33°C.
Temperature altered partitioning patterns in the whole plant
and at 33°C more leaves and less stem and roots were
produced, giving increased shoot to root ratios. Lamina leaf
folding also occurred at high air temperatures and the
authors implied that induced high vapor pressure deficits may
have been the cause. Severe constraints on data measurements
were imposed by the large nature of the banana plants and
restricted growth chamber space. This could, in part, explain
why critical physiological data such as photosynthesis,
transpiration and leaf water potential were not recorded.
Assimilate partitioning results were estimated using relative
growth rate formulae and no analyses on tissue carbohydrate
status were performed.
Root-Zone Temperature and Soil Moisture Interactions
The complexity of responses of RZT and soil moisture
stress alone are such that few researchers have attempted
studies on their interactive effects. Lai (80) reported in
1974 that a RZT of 35°C significantly decreased root and
shoot growth and transpiration in maize seedlings. Effects of
high RZT were aggravated by low soil moisture levels. Changes
in soil water supply and RZT were found to have predictable

19
effects on the relationship between leaf water potential and
transpiration rate in several tree species (37,54,68,135).
Kaufmann (69) evaluated the effects of RZT (6° to 20°C)
and two soil drying cycles (9 and 12 day) on water relations
and growth of Monterey pine (Pinus radiata. Don) under
greenhouse conditions. The slope of the relationship of xylem
pressure potential to transpiration was influenced by RZT.
Under well-watered conditions, neither transpiration nor
stomatal conductance were affected by RZT because xylem
pressure potential was not low enough to cause stomatal
closure. At the end of 9- and 12-day drying cycles, however,
transpiration was lower than in the well-watered controls as
a result of stomatal closure. RZT apparently had no consist¬
ent effect on transpiration rate. Reduced RZTs and soil
drying both significantly reduced shoot and root extension
after 28 days even though low RZT did not cause stomatal
closure.
In an earlier study with 'Monterey' pine, Babalola et
al. (5) measured photosynthesis, transpiration and
respiration at four soil water levels and four RZTs (10° to
27°C ). They found that the rates of photosynthesis, respi¬
ration and transpiration decreased with increasing soil water
deficits. However, only at soil water levels lower than -0.7
bars and at RZTs of 10°C and above were the rates of
photosynthesis and transpiration affected similar to changes
in stomatal conductance.

20
Barlow et al. (7) studied the effects of RZT and soil
water potential on corn seedlings. This investigation also
did not involve supraoptima1 temperatures but there were
notable interactive effects. Rates of leaf elongation, net
photosynthesis, transpiration and leaf water potential were
simultaneously monitored under varying stress conditions. In
their first study, leaf elongation was found to be much more
sensitive to changes in root temperature (12° to 28°C) than
either photosynthesis or transpiration and ceased at a LWP of
-0.9 MPa; physiological responses were not affected until
LWPs of -1.2 to -1.3 MPa were attained. A decrease in net
photosynthesis coincided with an increase in stomatal and
mesophyll resistance at low LWPs. The authors concluded by
emphasizing the need for whole plant studies so that stress
effects on photosynthesis, translocation and assimilate
partitioning could be better analyzed.
Two root temperatures (21.1° and 26.6°C) and two soil
water potentials (-0.35 and -2.50 bars) were employed in
their second experiment (8). Leaf elongation and total dry
matter were decreased 44% and 26%, respectively, soluble
carbohydrates increased 42% , while rates of transpiration
decreased 24% at the low soil water potential. There were
also some interactions between RZT and soil water content.
Dry weights for plants at 21.1°c and -0.35 bars were similar
to those at 26.6°C and -2.50 bars. Higher soluble
carbohydrate concentrations with increased water stress were
offset by increased RZT. Decreased total plant dry weight and

21
transpiration at lower soil water levels were slightly offset
by higher RZT, presumably through increased root conductance
and water absorption.
Most of the studies involving soil temperature and
moisture interactions on banana in the tropics have been
field oriented and have involved the use of soil mulches
which effectively reduce soil temperature and conserve soil
moisture (10,11). In most cases, mulching led to increased
WUE which was based on a yield to evapotranspirati on ratio
rather than on the physiological parameters of photosynthesis
and transpiration. In the case of coffee, there are
independent studies on soil temperature (39) and soil
moisture (4,72,138), but no reports on their interactive
effects

CHAPTER III
EFFECTS OF IRRIGATION VOLUME ON BANANA AND IXORA UNDER
TWO GROWING CONDITIONS
Introduction
Although 'Grande Naine' banana (Musa spp. AAA) is a
mesophytic, herbaceous plant requiring moist soils for
optimum production (127), it can be grown in marginal areas
where soil water is limiting. Irrigation scheduled when
available soil moisture depletion reaches 60% to 80% has been
shown to be optimum (10,11,31,116), but detailed
characterization of physiological responses in banana to
decreasing soil water levels is lacking.
Moisture stress is thought to be necessary for flower
induction (4) in coffee, a close relative of ixora (Ixora
chinensis L. 'Maui') within the Rubiaceae family (6), but
adequate irrigation (72) is essential for growth and mineral
absorption (138). Because of its importance as a major
beverage crop, there are several field (100,101) and
container studies (39,72,138,140) on the physiological
responses of coffee, but no work has been conducted on the
effects of varying soil water levels on container-grown
ixora. The present study investigated the physiological
responses of banana and ixora to three irrigation levels and
22

23
a 14-day drying cycle. Irrigation experiments were conducted
under both greenhouse and growth room conditions.
Materials and Methods
Plant Materials and General Cultural Procedures
Ten- to 12-cm tall tissue-cultured banana plants and
similar-sized, uniform ixora rooted cuttings were obtained
from commercial nurseries. Plants were hardened with
intermittent mist (6 sec min-1) in a glass greenhouse at 28°C
under 80% light exclusion for one week, then moved to 40%
light exclusion for another week. Plants were transplanted to
white 4-cm diameter x 21-cm tall conical containers (150 cm3)
using Metro-Mix 300 growth medium (W.R. Grace and Co.,
Cambridge, MA). Plants were then moved to the experimental
greenhouse or growth room where they were watered to
container capacity daily and allowed to acclimatize for one
week prior to the initiation of the experiment.
Greenhouse
Experiment 1. Experiment 1 was initiated in July, 1985,
in an air-conditioned greenhouse in which maximum
photosynthetic photon flux density (PPFD) ranged from 700 to
800 umol m-2 s-1, and temperatures were 25 to 30°C day and 18
to 21°C night. Relative humidity varied from 40 to 80%
(Figure 3-1).
Plants were watered daily at 2200 hr with 10+2 ml (Wl),
20+4 ml (W2) or 40 + 8 ml (W3) per container. These irrigation

I owrí) QJdd
100 O
0800 1000 1200 1400 1600
TIME (hrs)
Figure 3-1. Photosynthetic photon flux density and relative humidity in the
greenhouse during measurements of physiological parameters.
no
PERCENT RELflTIUE HUHIDlTYO

25
volumes provided 53% to 60%, 65% to 75%, and 85% to 100%
container capacity (CC) respectively, for banana and 52% to
64%, 68% to 80% and 90% to 100% CC, respectively, for ixora.
The comparatively smaller root mass of ixora was responsible
for the difference in growth medium waterholding capacities
between these plants. Water was applied automatically through
drip tubes by a battery-operated controller (Water Watch
Corp. Seattle, WA). One, two and four 1-mm drip
tubes/container constituted the Wl, W2 and W3 irrigation
treatments, respectively.
After 14 days of irrigation treatments, diurnal
measurements of leaf photosynthesis (PS), transpiration (TR)
and leaf conductance (CS) were made using a portable
photosynthesis system (Model LI-6000, LI-C0R, Inc., Lincoln,
NE). Measurements were initiated at 0800 hr EDT on a
cloudless day and taken every 2 hr until 1600 hr. A 1-liter
cuvette chamber was used for measurements and the mean of
eight consecutive 30-sec observations on each leaf
constituted a measurement. A zero check of the analyzer was
performed between treatments within each replicate.
Simultaneous measurements of LWP were made using a Scholander
type pressure chamber (PMS Instrument Model 600, Santa
Barbara, CA) as described by Barrett and Nell (9). The third
most recently expanded leaf in banana was selected for
measurements of both gas exchange and LWP. This leaf has been
shown to be the most responsive and the youngest with fully
developed stomata (127). Gas exchange processes in ixora were

26
measured on the most recently matured leaves but apical
shoots were sampled for LWP. Six plants per treatment were
sampled for all measurements.
Experimental design was a randomized complete block with
21 plants per water treatment within each of three blocks.
Six replicate plants were randomly selected at each sampling
time. Water use efficiency (WUE) was calculated from PS/TR.
Means and standard errors were calculated and plotted against
time of day.
Experiment 2. In order to further monitor physiological
responses of water stress and correlate these with visual
plant symptoms, water was withheld from banana and ixora
plants for 14 days beginning one week after transplanting.
This drying cycle study was conducted simultaneously with
experiment 1 in the same greenhouse and similar cultural
practices were used. Midday LWP and CS measurements were
recorded on five plants of each species at 2-day intervals
using the procedures described in experiment 1. A completely
randomized design was used in this experiment and means and
standard errors were calculated. There were five replicate
plants per treatment for each sampling date.
Growth room.
Experiment 3. Experiment 1 was repeated in a 3.0 m by
7.6 m walk-in growth room in order to investigate
physiological responses to water stress under more precisely
controlled environmental conditions. Irradiance of 1100 umol
m“2 s-1 PPFD, as measured by a quantum radiometer (LI-COR

27
Model LI-185A, LI-COR Inc., Lincoln, NE), was supplied at
plant canopy level by 1000 W phosphor-coated metal-arc HID
bulbs (GTE Sylvania Corp., Manchester NH) from 0600 to 1830
hr daily. Air temperatures of 28°C day and 21°C night and a
relative humidity of 65% to 70% were maintained. Three
irrigation volume treatments were applied as in experiment 1
in a randomized complete block design with three blocks and
21 plants per water treatment per block. Species were treated
and analyzed as separate experiments. On the fifteenth day
after water stress treatments, diurnal measurements were
initiated at 0800 hr and continued at 2-hr intervals until
1800 hr. Leaf sampling methods and measurement procedures for
PS, CS, TR and LWP were employed as in experiment 1.
Results and Discussion
Greenhouse
Experiment l--banana. Irrigation treatments influenced
daily PS, CS and TR maxima as well as diurnal patterns
(Figure 3-2A, 3-2B, 3-2C). Relatively high LWP were
maintained by plants in the W2 and W3 irrigation treatments
but there was a distinct decline in LWP by 1000 hr in plants
receiving the lowest irrigation volume, W1 (Figure 3-2D).
PS patterns of plants under the W3 and W2 irrigation
treatments paralleled the corresponding CS patterns, with a
rise in PS from 0800 hr to a peak at 1400 hr of 0.20 mg CO2
m“2 s“1 for the W3 treated plants. Maximum PS for plants in
the W2 irrigation treatment were reduced almost three-fold

Figure 3-2. Effects of three irrigation volumes on the diurnal physiological
responses of 'Grande Naine' banana grown under greenhouse conditions. A. leaf
photosynthesis B. leaf conductance C. transpiration and D. leaf water potential.
W1 = 10+2 ml, W2 = 20+4 ml and W3 = 40 +_8 ml daily per 150 cm3 container. Points
are the means of six replicate plants and vertical bars represent the SE.

0800 1000 1200 1400 1600 08 0 0 1000 1200 1400
TIME (hrs) TIME TRANSPFATTON (mg H20 m-2 s-1)
PHOTOSYNTHESIS (mflC02 m-2 s-1)
o o oi
O S fe 8 §
6Z

30
below plants at W3 (Figure 3-2A). PS for the W1 treated
plants was significantly reduced compared to plants in the
other irrigation treatments with a midday rate of only 0.003
mg CO2 m-2 s--*-.
CS in plants receiving the highest irrigation volume,
W3, averaged 0.37 cm s"1 at 0800 hr, had more than doubled by
1400 hr and declined sharply to 0.17 cm s-* at 1600 hr
(Figure 3-2B). CS in plants under the W2 irrigation followed
the same pattern, but was reduced by 50%, 47% and 49% at
1000, 1200 and 1400 hr, respectively. CS in plants at the
lowest irrigation level did not change appreciably throughout
the day and averaged only 0.08 cm s"l, suggesting that the
stomata remained nearly closed throughout the day.
Therefore, PS and CS were severely reduced by decreased
irrigation and both declined in parallel.
Although TR increased from 0800 hr to 1000 hr for both
W2 and W3 treated plants, TR in W2 and W1 plants was
significantly lower (Figure 3-2C). TR remained somewhat
stable between 1000 and 1400 hr for all treatments and
declined significantly between 1400 and 1600 hr under the W3
irrigation volume. Maximum TR was 44.4, 14.2 and 6.3 mg H2O
m-2 s-1 for the W3, W2 and W1 treated plants, respectively.
LWP of plants under the W2 and W3 irrigation treatments
were fairly stable throughout the day, but LWP of plants at
the lowest irrigation level declined significantly after 0800
hr to a low LWP of -0.90 MPa at 1000 and 1200 hr (Figure 3-
2D). Although there was an increase in LWP for W1 treated

31
plants after midday, the LWP at 1600 hr of -0.62 MPa remained
significantly lower than the 0800 hr LWP. The reduction in CS
and the decline in gas exchange processes but not leaf water
status at midday and 1400 hr for the W2 treated plants,
indicated that there were factors other than leaf water
status controlling stomatal opening.
Water use efficiency (WUE) for plants under the W3 and
W2 irrigation levels increased almost linearly throughout the
day but did not differ between treatments until 1200 hr,
after which WUE in W2 treated plants was higher (Figure 3-3).
This indicated that TR in plants under the W2 treatment was
reduced more relative to PS. However, the WUE pattern for the
severely stressed W1 treated plants remained low throughout
the day. The leaf folding mechanism commonly observed in
field-grown banana plants at midday hours (127,136) was
observed in the W2 treated plants. Increased WUE values,
though calculated on a yield/evapotranspiration ratio, have
also been obtained for field-grown bananas under moderate
reductions in soil moisture levels (11).
Altered physiological processes in banana with
decreasing irrigation levels agrees with Shmueli's work (126)
in which TR and CS in Cavendish banana were reduced by
available soil moisture levels below 66%. Chen (25) also
found a similar effect at a soil water level of 50% to 60%
field capacity. These were both field studies in which soil
moisture effects could have been confounded by ambient vapor

Figure 3-3. Effects of three irrigation volumes on the diurnal water use
efficiency of 'Grande Naine' banana grown under greenhouse conditions. W1
ml, W2 = 20+4 ml and W3 = 40jj8 ml daily per 150 cm^ container. Points are
means of six replicate plants and vertical bars represent the SE.
= 10 + 2
the

0800 1000 1200 1400 1600
UflTER USE EFFICIENCY (mg C02/mg H2O.10-4)
u) cn cd
q Q Q Q
ee
120

34
pressure deficits to a greater extent than occurred in the
present greenhouse experiment. CS and TR reported for both
field studies, however, were closely correlated. Brun (20)
reported good correlation between PS and TR in intact banana
leaves in response to light intensity but CS was not
monitored in his study. Results in the present experiment
indicate that stomatal closure was associated with reductions
in PS and TR.
Severe growth reduction and leaf chlorosis have been
induced by soil moisture depletion levels below 50% in field
studies (126). These effects were generally attributed to
reductions in PS, although PS rates were not measured. The
present study substantiates these hypotheses, since the
lowest irrigation level produced similar water stress
symptoms and significantly decreased PS (Figure 3-2A).
Experiment l--ixora. Although irrigation treatments
highly influenced measured parameters, diurnal patterns did
not necessarily parallel one another (Figure 3-4). Under the
W3 irrigation treatment, maximum PS and TR rates of 0.26 mg
CO2 m-2 s"1 and 58.1 mg H2O m-2 s-*, respectively, occurred
at 1400 hr (Figure 3-4A, 3-4C). However, CS was somewhat
erratic, with a maximum at 1000 hr, followed by a decline at
1200 hr and a small rise at 1400 hr (Figure 3-4B). PS and CS
were significantly reduced by the W2 and W1 irrigation levels
through 1400 hr compared to W3 and under the W1 level CS and
TR appeared to be more closely related.

Figure 3-4. Effects of three irrigation volumes on the diurnal physiological
responses of Ixora chinensis L. 'Maui' grown under greenhouse conditions. A. leaf
photosynthesis 6. leaf conductance C. transpiration and D. leaf water potential.
W1 = 10+_2 ml, W2 = 20+4 ml and W3 = 40+jB ml daily per 150 cm3 container. Points
are the means of six replicate plants and vertical bars represent the SE.

0800 1000 1200 1400 1600 ¿íaÓO 1000 1200
TIME (hrs) TIME (hrs)
TRANSPFATON (mo H20 m-2 s-1) PHOTOSYNTHESIS (mg 002 m-2 *-1)
9e

37
LWP was generally reduced by decreasing irrigation
volumes (Figure 3-4D). Midday LWP for the W2 and W1
treatments were 2 and 2.5 times less than for plants under
the W3 irrigation treatment, respectively. Irrigation
treatments, therefore, were well distinguished by measuring
LWP.
Reduced irrigation, however, decreased PS relatively
more than TR as reflected in the higher WUE in plants at the
W3 irrigation level compared to plants under the W2 and W1
levels (Figure 3-5). Non-stoma tal inhibition of PS under
water stress conditions have also been reported in other
species (14,15) and may have contributed to the lower WUE
observed with water stress in this study.
Although midday declines in CS have been noted for
coffee (100,101,140) under well-watered field conditions, the
suggestion that light was responsible (101) would not seem to
apply to ixora. In the high light environment of the growth
room CS did not decline at midday (Figure 3-10B).
A high irrigation treatment of 200 ml/15 cm pot/day
induced flooding effects and caused leaf abscission in coffee
(72). Such effects were not induced in this study by the
highest irrigation treatment, W3, but the erratic midday
stomatal response referred to above might have been in
response to the high watering level.

Figure 3-5. Effects of three irrigation volumes on the diurnal water use
efficiency of Ixora chinensis L. tMaui ' grown under greenhouse conditions. W1
10 + 2 ml, W2 = 2 0 + 4 ml and W3 = 40 + 8 ml daily per 150 cm^ container. Points are
the" means of si x”repl i cate plants“and vertical bars represent S.E.

UIRTER USE EFFICIENCY (mg C02/,,n9
&
J
o
•—*
d
CM
i
60
45
15
0
Ull
U2
0800
1000 1200
1400
1600
TIDE (hrs)
co
co

40
Experiment 2--banana. LWP decreased linearly while CS
decreased in a sigmoidal pattern during the 14 days without
irrigation (Figure 3-6). Leaf folding along the midrib is a
common feature of water-stressed field-grown banana plants at
midday hours (127,136), but plant water status associated
with the mechanism has not been previously defined. In this
study, leaf folding occurred between day 4 and 6 and
coincided with a decline in LWP from -0.51 to -0.65 MPa and a
decrease in CS from 0.46 cm s"l on day 4 to 0.051 cm s"* on
day 6. In experiment 1, leaf folding was also observed in the
W2 treated plants at midday with comparable LWP and CS. This
response therefore, substantiated the irrigation-volume
method as a fairly accurate, practical means of imposing
water stress treatments in banana. Anatomical studies have
indicated a special band of cells along the midrib of the
banana leaf (130) which are responsible for the folding
response and it is likely that these cells are more
responsive to decreasing LWP than the other leaf cells.
Further reduction in LWP to -0.91 MPa and CS to 0.019 cm
s_1 probably impaired the PS mechanism since chlorosis
occurred indicating greater chlorophyll degradation than
synthesis (57). At day 14, when LWP declined to -1.34 MPa and
CS to 0.004 cm s"l, there was severe chlorosis and apparent
growth cessation. At this stage leaves remained folded
probably indicating a cessation in gas exchange processes and
complete stomatal closure.

o
Leaf Folding
Miid Chkxosé
.80'
DAYS AFTER WATERNG
Figure 3-6. Effect of a 14 day drying cycle on leaf water potential and leaf
conductance of 'Grande Naine' banana grown under greenhouse conditions. Points
are the means of five replicate plants and vertical bars represent the SE.

42
Experiment 2--ixora. LWP decreased linearly after day 2
but there was a steep decline in CS from 0.53 cm s_1 on day 2
to 0.14 cm s_1 on day 4 (Figure 3-7). Wilting of youngest
leaves occurred between day 4 and 6 and corresponded to
decreases in LWP from -1.08 to -1.71 MPa and in CS from 0.14
to 0.038 cm s"1. Severe wilting of top leaves and abscission
of bottom leaves were observed on day 8 at a LWP of -2.06 MPa
and a CS of 0.008 cm s~l. Plant water status between days 4
and 6 of the cycle was comparable to that at midday
(-1.2 MPa) in plants under the W1 irrigation treatment in
experiment 1. This indicated that reducing irrigation volume
was an effective means of imposing water stress in ixora.
Beyond the day 8 stage, wilting and leaf abscision made it
difficult to record further data. Since stressed plants were
not rewatered it was impossible to determine whether stomatal
closure and leaf abscission observed in this study were
clearly mechanisms imparting stress tolerance that allowed
the plants to survive.
Growth Room
Experiment 3--banana. PS, CS and TR diurnal patterns
paralleled one another under the more controlled conditions
of the growth room (Figure 3-8) but irrigation treatment
effects were not quite as dramatic as in the greenhouse
experiment (Figure 3-2). Maximum PS, CS and TR occurred in
the growth room at 1000 hr followed by a gradual decline to

Figure 3-7. Effects of a 14 day drying cycle on leaf water potential and leaf
conductance of Ixora chi nensi s L. 'Maui' grown under greenhouse conditions.
Points are the means of five replicate plants and vertical bars represent the SE.

Figure 3-8. Effects of three irrigation volumes on the diurnal physiological
responses of 'Grande Naine' banana grown under growth room conditions. A. leaf
photosynthesis B. leaf conductance C. transpiration and D. leaf water potential.
W1 = 10+2 ml, W2 = 20+4 ml and W3 = 40+8 ml daily per 150 cm^ container. Points
are the means of six replicate plants and vertical bars represent the SE.

TRANSPIRATION (mg H20 nrr-2 8-1) PHOTOSYNTHESIS (mg C02 m-2 s-2]

46
1800 hr (Figure 3-8) while maximum levels in the greenhouse
experiment were at 1200 to 1400 hr for CS and 1000 to 1400 hr
for TR. Higher light intensity early in the day in the growth
room could have resulted in earlier stomatal opening (33).
Both Shmueli (126) and Chen (25) reported CS and TR diurnal
maxima in banana at 1000 hr followed by declines and with
minor increases at 1400 hr.
Under the W3 irrigation treatment, maximum PS, CS and TR
were 0.596 mg CO2 m“2 s"*, 0.720 cm s“* and 89.2
mg H2O m-2 s-1, respectively at 1000 hr. PS, CS and TR were
reduced 252, 482 and 352 at 1000 hr, respectively, by the W2
irrigation treatment compared to the W3 treatment (Figure 3-
8A, 3-8B, 3-8C). At the lowest irrigation level, all gas
exchange processes remained consistently low throughout the
day and maximum PS, CS and TR were 0.097 mg CO2 m-2 s"*,
0.181 cm s"1, and 38.2 mg H2O m-2 s“*, respectively.
LWP generally declined for all irrigation treatments
after 0800 hr for most of the day with an increase after 1400
hr for the W3 treated plants (Figure 3-8D). Midday LWPs were
-0.58 MPa, -0.66 MPa and -0.82 MPa for the W3, W2 and W1
treatments, respectively. LWP between 1000 hr and 1400 hr
were not different for the W3 and W2 treatments but LWP was
significantly reduced by the W1 irrigation level. Despite the
similar LWP between W2 and W3 at 1000 to 1400 hr, significant
differences in PS, CS and TR were observed.
Davies (33) reported increased stomatal sensitivity to
decreasing soil water levels for cotton and soybean under

47
growth chamber conditions but Boyer (14) found PS of
sunflower to be affected more by photochemical activity than
by stomatal conductance under similar conditions. In the
present study there were relative increases in PS but not CS
in plants grown in the growth room as compared to the
greenhouse.
Maximum CS and TR have been shown to occur in moderately
watered field-grown banana at 1000 hr with subsequent
declines and smaller peaks at 1400 hr (25,126). It is notable
that this response resembled the diurnal patterns that were
observed in the W2 treated plants in the present study. In
the growth room LWP were generally lower than in the
greenhouse and could have been responsible for the general
decline in PS, CS and TR after 1000 hr.
Under growth room conditions, the W3 irrigation
treatment generally maintained a constant WUE throughout the
day (Figure 3-9) reflecting the parallel changes in PS and
TR. Though WUE for the W2 treated plants declined at 1200 and
1400 hr compared to 0800 hr, it was still not significantly
different from that of the W3 plants during these hours. The
relative improvement in WUE at the lowest irrigation in this
experiment compared to the same treatment in the greenhouse
study was due mainly to increased PS. The W1 irrigation
treatment under growth room conditions therefore did not
depress physiological responses as much as that observed in
the greenhouse study.

Figure 3-9. Effects of three irrigation volumes on the diurnal water use
efficiency of 'Grande Naine' banana grown under growth room conditions. W1 = 10+_2
ml, W2 = 20+4 ml and W3 = 40+8 ml daily per 150 cm^ container. Points are the
means of six replicate plants and vertical bars represent the SE.

I
o
o
(N
X
cn
E
\
(N
O
U
cn
E
>*
U
z
u
LJ
UJ
in
3
X
X
3
120 -
80 --
40 --
0
0800
1000
1200 1400 1 BOO 1800
TinE (hrsJ
-p*

50
Experiment 3--ixora. Linder growth room conditions, the
W2 and W1 treatments did not consistently decrease
physiological parameters (Figure 3-10) as severely as in the
greenhouse study (Figure 3-4). At 1200 hr, there were no
differences in responses of plants under the W2 and W3
treatments but at the lowest irrigation treatment, all
parameters were significantly reduced over those at the
higher irrigation levels.
Maximum PS rates were 0.425, 0.388 and 0.151 mg CO2 m~2
s"l for the W3, W2 and W1 irrigation treatments, respectively
(Figure 3-10A). These rates were 1.25, 7 and 10 times higher
than the respective rates in the greenhouse study and may
have reflected the influence of increased PPFD on PS in
ixora, particularly at the lower irrigation treatments. Light
levels in the greenhouse were probably below light saturation
levels for ixora.
There were no differences in CS between plants at the W2
and W3 irrigation levels from 0800 to 1400 hr but CS was
significantly reduced at these hours by the W1 irrigation
volume (Figure 3-10B). Midday CSs were 0.492, 0.382 and 0.190
cm s-1 under the W3, W2 and W1 treatments, respectively.
Compared with CS in plants in the greenhouse experiment
(Figure 3-4B), these values were 342, 492 and 872 higher at
the W3, W2 and W1 irrigation levels, respectively. The
increased response in stomatal opening, especially at the
lower irrigation levels corresponded with increases in PS.

Figure 3-10. Effects of three irrigation volumes on the diurnal physiological
responses of Ixora chinensis L. 'Maui1 grown under growth room conditions. A.
leaf photosynthesis B. leaf conductance and C. transpiration. W1 = 10+2 ml, W2 =
20+4 ml and W3 = 40+8 ml daily per 150 cm^ container. Points are the means of six
replicate plants ancf vertical bars represent the SE.

o
0»
o
o
o.
o
m
00
PHOTOSYNTHES8 (mg 002 m-2 »-1)
$ fe ,1
TRANSPRAT10N (mg H20 m-2 a-1)
o P 8 8
m
?
9
O
00
o
o
2
m —'
9
LEAF CONDUCTANCE (cm a-1)
°—& 11
3S

53
The high PS rate at 0800 hr under the W3 irrigation
resulted in a correspondingly high WUE (Figure 3-11) and
although WUE then declined at 1000 and 1200 hr for the
highest irrigation treatment, WUE values were basically the
same for all irrigation treatments through the rest of the
day. Under greenhouse conditions, W3 treated plants clearly
had higher WUE than plants subjected to lower irrigation
levels (Figure 3-5). The increase in WUE in the W2 and W3
relative to the W3 treated plants under growth room
conditions was due mainly to the increased PS rates in the
high light environment.
Growth room conditions, therefore, not only moderated
physiological responses of ixora to reduced irrigation volume
but also stabilized PS/TR changes as reflected by WUE values.
The increased photosynthetic response in plants grown under
the W1 and W2 irrigation volumes suggested that ixora could
probably better withstand decreased irrigation volume in the
growth room compared to greenhouse conditions reported in
this study.

Figure 3-11. Effects of three irrigation volumes on the diurnal water use
efficiency of Ixora chinensis L. 'Maui' grown under growth room conditions. W1
10 + 2 ml, W2 = 20 + 4 ml and W3 = 40+^8 ml daily per 150 cm3 container. Points are
the means of six replicate plants and vertical bars represent the SE.

UlflTER USE EFFICIENCY (mg C02/mg
i
o
o
(N
I
80
40
0
I 1 1 1 1 h-
0800 1000 1200 1400 1600 1800
TinE (hrs )
cn
tn

CHAPTER IV
ROOT-ZONE TEMPERATURE EFFECTS ON BANANA AND IXORA
UNDER TWO GROWING CONDITIONS
Introduction
High root-zone temperatures (RZT) have been shown to
reduce plant growth (59,63,70) and affect many physiological
processes (43,46,64). Tropical and subtropical plant species
are often thought to be heat tolerant, but soil temperatures
as high as 52°C have been recorded in the tropics (39), and
temperatures in this range can be lethal for some tropical
(39,63) and subtropical crops (62). An air temperature of
33°C was reported as being optimum for growth and dry weight
partitioning in banana (Musa spp. AAA) (142) and mineral
composition was highly influenced by temperatures from 18 to
33°C (143). A RZT of 33°C was found to be optimum for growth
and transpiration in coffee (39), a member of the same
subfamily of the Rubiaceae to which ixora belongs (6).
However, there are no reports on RZT effects on growth and
physiology of container-grown banana or ixora.
It is essential to identify growth and physiological
effects of high RZT in container production systems before
control measures can be developed to alleviate such effects.
This study investigated the short-term effects of RZT on
56

57
container-grown 'Grande Naine' banana and Ixora chinensis L.
'Maui' under greenhouse and growth room conditions.
Materials and Methods
Plant Materials and General Cultural Procedures
Ten- to 12-cm tissue-cultured 'Grande Naine' banana
plants and similar-sized, uniform I xora c hin e n sis L. 'Maui*
rooted cuttings were obtained from commercial nurseries.
Plants were hardened in intermittent mist (6 sec min"1) for
one week under 80% light exclusion and then moved to 40%
light exclusion for another week. Plants were then
transplanted to 4-cm diameter x 21-cm high conical containers
(150 cm^) using Metro-Mix 300 growth medium (W.R. Grace and
Co., Cambridge, MA). Plants were fertilized weekly with a
soluble 20N-8.8P-16.6K fertilizer (Peters 20-20-20, W.R.
Grace and Co., Cambridge, MA) at 150 ppm N.
Experiment 1--Greenhouse
An experiment was initiated August, 1985 in an air-
conditioned glass greenhouse with a mean maximum PPFD of 600
to 700 umol m"2 s-* PPFD, and a 25° to 30°C day and 18° to
21°C night temperature. Relative humidity was not controlled
and varied from 40% to 80% (Figure 4-1). Recently
transplanted banana and ixora plants were watered daily to
container capacity and allowed to acclimatize for one week.
RZT treatments were established within styrofoam-lined
wooden air bath boxes (1 m x 1 m x 20 cm) in which plant
containers were firmly inserted to within 2 cm from their

I ornrí) Qjdd
100
O
x
1000
X
~ 800
i
Ul
CM
E
600
400
200
0
80
60
40
20
0800 1000 1200 1400 1600
TJflE (hrs 1
Figure 4-1. Photosynthetic photon flux density and relative humidity in the
greenhouse during measurements of physiological parameters.
PERCENT RELñTUdE HUniDITY O

59
upper rims. Four equally spaced 100 watt aluminum foil-
covered incandescent light bulbs provided convective heating
of the enclosed root systems. Proper distribution of heat and
aeration within the boxes were ensured by two fans (IMC
Magnetics, Roch. NH) per box. The light bulbs in each box
were controlled by a pre-set thermostat and the temperatures
in each box were verified daily with a thermocouple
thermometer (TH 65, Wescor, Inc., Logan, UT). Electrical
power to the temperature boxes was controlled by Intermatic
Time Controls (Intermatic Inc. Spring Grove, Ill), providing
daily RZT treatments from 1000 to 1600 hr. Boxes were placed
on 1 m high benches and an automatic drip irrigation system
provided daily watering of all plants to container capacity
at 2200 hr.
RZT treatments established were 28^1°, 33+_l°, 38^1° and
43+_l°C. Each temperature box contained six ixora and six
banana plants and there were three replicate boxes for each
treatment temperature arranged in a randomized complete block
design. Species were treated and analyzed as separate
experiments. Means and standard errors were calculated for
the six replicate plants measured at each sampling time.
After 14 days of RZT treatments, diurnal measurements of
leaf photosynthesis (PS), leaf conductance (CS),
transpiration (TR) and leaf water potential (LWP) were made
using a portable photosynthesis system (LI-6000 model, LI¬
COR, Inc., Lincoln, NE). Measurements were initiated at 0800
hr EDT on a cloudless day and taken every 2 hr until 1600 hr.

60
A 1-liter cuvette was used and the mean of eight consecutive
30-sec observations constituted a measurement. A zero check
of the analyzer was performed between treatments within each
replicate. Simultaneous measurements of LWP were made using a
pressure chamber (PMS Instrument Model 600, Santa Barbara,
CA) as described by Barrett and Nell (9). The third most
recently expanded leaf in banana was selected for both
gaseous exchange processes and LWP measurements. This leaf
has been shown to be the most responsive and the youngest
with fully developed stomata (127). The most recently matured
ixora leaves were used for PS, CS and TR measurements and 4-
to 5-cm shoot tips were sampled for LWP.
Experiment 2--Growth Room
In order to further monitor the physiological responses
of ixora and 'Grande Naine' banana to RZT under more
precisely controlled environmental conditions, experiment 1
was repeated in a 3.0-m by 7.6-m walk-in growth room. Light
irradiance of 1100 umol m-2 s-* PPFD, as measured by a
quantum radiometer (LI-C0R Model LI-185A, LI-COR Inc.,
Lincoln, NE), was supplied from 0600 to 1830 hr at plant
canopy height by 1000 W phosphor-coated metal-arc HID bulbs
(GTE Sylvania Corp., Manchester, NH). Air temperature of 28°C
day and 21°C night and a relative humidity of 65% to 70% were
maintained.
Each conical container with a transplant was suspended
through a tightly fitting styrofoam ring within a specially

61
constructed root heating tube (RHT). Each RHT was 22.5 cm
high and constructed from 7.5 cm diameter metal pipe wrapped
with 60 watt 120 vac heating tape (Smith-Gates Corp.,
Farmington, CT) and 1.25 cm thick foam insulation. The RHTs
were connected to solid state electronic controllers which
maintained preset treatment temperatures by a thermistor
feedback mechanism. Each controller maintained treatment
temperatures in 16 tubes with four tubes at each of four
specified temperatures. Treatment temperatures were set and
also maintained at 28+0.3°, 33+0.3°, 38+0.3° and 43+0.3°C.
Similar leaf number and sampling methods were used and the
same physiological measurements were recorded as in
experiment 1 after 14 days of root-zone temperature
treatments.
Experimental design was a randomized complete block with
36 plants of each species per temperature treatment arranged
in three blocks. Species were treated and analyzed as
separate experiments. Means and standard errors were
calculated from six replicate plants measured at each
sampling time.
Results and Discussion
Experiment 1--Greenhouse
Banana. At the 33° and 28°C RZTs, PS increased to midday
maxima of 0.53 and 0.50 mg C 0 2 m~^ $ — 1 ^ respectively, and
then declined (Figure 4-2A). Maximum PS was 0.43 mg CO2 m“2
s“l at 1000 hr in the 43°C RZT treated plants, while PS was

Figure 4-2. Effects of four root-zone temperatures on the diurnal physiological
responses of 'Grande Naine' banana grown under greenhouse conditions. A. leaf
photosynthesis B. leaf conductance C. transpiration and D. leaf water
potential. Points are the means of six replicate plants and vertical bars
represent the SE.

£9

64
0.31 mg CO2 m~2 S-1 f0r plants at the 38°C RZT. These rates
were 202 and 422, respectively, of the maximum PS recorded at
33°C.
Midday CS for the 28° and 33°C RZT treated plants were
0.43 and 0.34 cm s*1, respectively, with significant
reductions to 0.28 and 0.19 cm s~l for plants at the 38° and
43°C RZT (Figure 4-2B). TR diurnal patterns were closely
related to those for CS, with maximum rates of 54 and 52 mg
H2O m-2 s-1 at midday for plants at the 28° and 33°C RZT,
respectively (Figure 4-2C). Midday TR was significantly
reduced by the 38° and 43°C RZT compared to the other RZTs.
LWPs were generally not different throughout the day for
the 28°, 33° and 38°C RZT treated plants. Among these three
RZT treatments, a low midday LWP of -0.52 MPa was recorded at
the 38°C RZT (Figure 4-2D). A significant reduction in LWP
was noted for plants at the 43°C RZT at 1200 hr, with a
midday LWP of -0.76 MPa. Although transpirational water loss
was reduced in plants at the 43°C RZT at midday compared to
plants at the 28° and 33°C RZTs, LWP of plants at the highest
RZT were significantly lower than those of the other three
RZT treatments. At 38°C, TR was less than at 28° and 33°C RZT
but LWPs were the same between these treatments. These
responses may suggest impairment of the absorptive and/or
conductive capacity of roots in plants exposed to the 43°C
RZT.
In summary, PS in banana plants was greatest at the 28°
and 33°C RZTs under greenhouse conditions. The 28°, 33° and

65
38°C RZT treatments resulted in the highest LWP and the 43°C
RZT reduced all measured parameters. RZTs of 28° and 33°C are
well within the range of RZTs reported as being optimum for
the growth of coffee (39), tomato (42) and pepper (44) while
38°C RZT has reportedly decreased root conductance (146) and
transpiration (73).
Ixora. PS in plants at all RZT treatments increased from
0800 hr to midday (Figure 4-3) but PS, CS and TR were
greatest at midday in plants exposed to the 33°C RZT. LWP
declined for most treatments at 1200 hr, with progressively
lower values from the 28° and 33°C RZT to the 38° and 43°C
treatments (Figure 4-3D).
Maximum midday PS rate was 0.41 mg CO2 m-2 s-1 for the
33°C treated plants with reductions to 0.29, 0.28 and 0.25 mg
CO2 m-2 s"1 for the 38°, 43° and 28°C RZT treatments,
respectively (Figure 4-3A). At 1400 hr, however, PS rates at
the 43° and 38°C RZT were 0.26 and 0.23 mg CO2 m-2 s“•*■,
respectively, while PS at 33°C RZT had declined to 0.17 mg
CO2 m-2 s-l. At 1600 hr, PS rates for the 28°, 33° and 38°C
RZT treated plants were not different. The 33°C RZT clearly
resulted in the highest midday PS but because this RZT did
not alter morning rates and caused rapid post-midday
declines, mean daily PS at 33°C was probably similar to that
at other RZT treatments.
CS and TR diurnal patterns were closely related, with
the midday maxima of 0.53 cm s-1 and 95 mg H2o m“2 s_1,
respectively, for both parameters occurring in the 33°C RZT

Figure 4-3. Effects of four root-zone temperatures on the diurnal physiological
responses of Ixora chinen sis L. 'Maui' grown under greenhouse conditions. A. leaf
photosynthesis BT leaf conductance C. transpirati on and D. leaf water
potential. Points are the means of six replicate plants and vertical bars
represent the SE.

0 8 0 0 1000 1200 1400 1600 08 0 0 1000 1200
TIME (hrs) TIME (hrs)
TRANSPRATTON (mg H20 m-2 8-1)
o 8 § 8 §
i i ■ —
PHOTOSYNTHESIS (mg C02 m-2 8-1)
L 9

68
plants (Figure 4-3B, 4-3C). Midday CS and TR were
significantly reduced in plants at the 43°C RZT compared to
the 33°C RZT but the 38°C RZT resulted in the lowest midday
TR and CS.
Mean midday LWP for plants at the 28° and 33°C RZT was
-0.71 MPa with progressive reductions by the 38°C (-0.87 MPa)
and 43°C (-1.02 MPa) RZT treatments (Figure 4-3D).
The 33°C RZT treated ixora plants exhibited highest
midday PS, CS, TR and had relatively good plant water status
compared with other RZT. Although PS was not measured, Franco
(39) reported that a RZT of 33°C was optimum for coffee in
terms of TR and root and shoot dry weights. TR and plant dry
weights were reduced with increasing RZT above 33°C and
plants died at 48°C RZT.
Midday LWPs decreased significantly to -0.87 MPa by the
38°C RZT treatment and to -1.02 MPa by the 43°C RZT
treatment, but visible symptoms of water stress were never
apparent in these plants. In contrast, water-stressed ixora
plants (experiment 2, chapter III) exhibited wilting around a
LWP of -1.08 MPa. Therefore, there may have been mechanisms
to maintain turgor under decreasing LWP in plants at the 38°
and 43°C RZT or plants may have become conditioned to the
high RZTs.
Experiment 2--Growth Room
Banana. Plants held at the 38°C RZT under growth room
conditions (Figure 4-4A) had a maximum midday PS rate of 0.81
mg CO2 m-2 s“■*•, which was a 54% increase over the

Figure 4-4. Effects of four root-zone temperatures on the diurnal physiological
responses of 'Grande Naine' banana grown under growth room conditions. A. leaf
photosynthesis B. leaf conductance C. transpiration and D. leaf water
potential. Points are the means of six replicate plants and vertical bars
represent the SE.

A
w 1jO
2 8oC
33oC
38oC
43oC
IMIIUIIIUIUUU
* » i ►
08 0 0 1000 1200 1400 1600 1800
LEAF WATER PO
0
L t * ■ 4 4—- I '*•
0800 1000 1200 1400 1600 1800
TIME (hrs)
'-si
o

71
corresponding greenhouse rate (Figure 4-2A). The 33°C RZT
induced a rate of 0.74 mg CO2 m-2 s-1 at 1000 hr while the
43°C RZT treated plants had the lowest PS rate of 0.39 mg CO2
m-2 s_1 at 0800 hr and through the afternoon hours(Figure 4-
4A).
Maximum stomatal opening for the 28° and 33°C RZT
occurred earlier in the day than for the higher RZT
treatments (Figure 4-4B). While maximum CS of 0.72 and 0.63
cm s"l occurred in plants at 1000 hr under the 28° and 33°C
RZT respectively, peak values for the 43° and 38°C RZT were
0.52 at midday and 0.70 cm s-1 at 1400 hr, respectively
(Figure 4-4B). The 43°C RZT caused significant reductions in
CS and TR from 0800 hr to 1000 hr and later in the day at
1800 hr. Highest TR rates occurred with the 38°C RZT with
values of 94.2 and 92.7 mg H20 m-2 s"1 at 1000 and 1400 hr,
respectively (Figure 4-4C).
Diurnal patterns of LWP were significantly influenced by
RZT treatments (Figure 4-4D). The 28°C RZT treated plants had
the highest midday LWP of -0.55 MPa, which was higher than
the LWP at the 33° and 38°C RZT treatments. The lowest midday
LWP of -1.26 MPa was induced by the 43°C RZT. Overall, LWP
were lower in plants in this experiment than those observed
in the greenhouse environment. This could be attributed to
the higher CS and TR rates that were exhibited by plants in
the growth room experiment compared to the greenhouse. In
both environmental conditions, however, the highest RZT of
43°C induced the lowest LWP with higher LWP at lower RZTs.

72
Although the 38° and 43°C RZTs appeared to induce water
stress in banana as reflected by low LWP, there was no leaf
folding and/or chlorosis as shown by plants that had been
water stressed to -0.52 and -0.66 MPa by irrigation
treatments under greenhouse and growth room conditions,
respectively (experiments 1,3 Chapter III).
Under growth room conditions, the 33° and 38°C RZTs
induced the highest PS rates though at different times of the
day. Under sunlit growth room conditions, 33°C air
temperature was reported as being optimum for growth and
partitioning in Cavendish banana but 37°C caused leaf injury
(142). Since growth measurements were not recorded in this
study it was impossible to determine whether the increased PS
observed at 38°C RZT resulted in increased growth or was
merely a response to the increased root sink demand for
photosynthates. The response of the banana plants grown at
38°C RZT in the two environmental conditions would indicate a
greater tolerance to RZTs up to 38°C under growth room
conditions. The 43°C treatment was supraoptimal under both
environments.
Ixora. Maximum PS, CS and TR at midday followed by
distinct declines were not as evident in ixora plants under
growth room conditions (Figure 4-5) as they were in the
greenhouse study (Figure 4-3). In plants subjected to the 28°
and 33°C RZT treatments, PS rates of 0.51 and 0.47 mg CO^ m“2
s“l, respectively, were recorded at 1000 hr and followed by a
gradual decline to 0.21 and 0.12 mg CO2 m-^ s_1,

Figure 4-5. Effects of four root-zone temperatures on the diurnal physiological
responses of Ixora chinenesis L. 'Maui' grown under growth room conditions. A.
leaf photosynthesis fH leaf conductance and C. transpiration. Points are the
means of six replicate plants and vertical bars represent the SE.

0800 1000 1200
PHOTOSYNTHESIS (mg C02 m-2 8-1)
&&

75
respectively, at 1800 hr (Figure 4-5A). A PS rate of 0.57 mg
CO2 m“2 S-1 Was observed at 0800 hr in the 38°C RZT treated
plants and was followed by a significant decline at 1000 hr
with no further decreases for the rest of the day. PS rates
were reduced during the morning hours in plants treated with
the 43°C RZT relative to other treatments but increased to
levels comparable to other RZTs at 1400 and 1600 hr. Except
for shifts in time at which maximum daily PS occurred, there
appeared to be little differences in overall PS values
between RZT treatments.
Diurnal CS (Figure 4-5B) and TR (Figure 4-5C) followed
the same general pattern. Maximum values for both parameters
occurred at 1200 and/or 1400 hr in plants treated with either
the 28°, 33° or 38°C RZT, with subsequent declines for the
rest of the day. Maximum CS recorded for these RZTs was
0.51 cm s“l at 1200 hr which was comparable to the
corresponding midday CS in the greenhouse. However, a CS of
0.46 cm s~l at 1400 hr in the growth room experiment
represented a 302 increase over the CS recorded at this hour
in the greenhouse. A similar pattern was observed for TR.
Maximum TR was 83.3 and 80.1 mg H2O m"2 s"* at 1200 and
1400 hr, respectively. CS and TR were significantly reduced
throughout most of the day in the 43°C RZT treated plants
(Figure 4-5B, 4-5C).
Under growth room conditions, therefore, RZTs of 28°,
33° and 38°C did not produce appreciable differences in
physiological responses in ixora. The relatively low PS, CS

76
and TR early in the day with subsequent increases at later
hours in the 43°C treated plants suggested a lag effect of
this RZT on stomata! opening.
Midday CS and TR were comparable to those observed in
the greenhouse. However, higher values for these parameters
were recorded at 1400 hr and later in the growth room as
compared to the distinct post-midday declines in the
greenhouse. There was an apparent effect of the constant high
light intensity during the illumination period in the growth
room that maintained midday stomatal opening to later hours.
However, the eventual decline in gas exchange processes at
1800 hr in the growth room suggested that the stomata may not
have completely lost the diurnal rhythmic pattern evident
under greenhouse conditions.

CHAPTER V
ROOT-ZONE TEMPERATURE AND SOIL MOISTURE EFFECTS ON
BANANA AND IXORA UNDER TWO GROWING CONDITIONS
Introduction
High temperature and water stress often occur
simultaneously in crop situations and alleviating the latter
is often considered a remedy for the former. Although many
drought tolerant plants are also heat tolerant (85),
interactions and sometimes negative correlations occur
between heat and drought stress (102). Research has tended to
concentrate on water and temperature stress effects
separately with little emphasis on defining or separating the
characteristics of these two stress factors.
Suboptimal root-zone temperature (RZT) (7,8,18) and
temperate plant species (5,18,69) have been employed in most
of the reported investigations that involve RZT and soil
moisture interactions. Barlow (7) found that increased shoot
carbohydrate under reduced soil moisture was offset by
increasing RZT in corn seedlings. Kaufmann (69) found that
soil water deficits directly affected leaf water potential
(LWP) and transpiration (TR) in 'Monterey' pine seedlings but
RZT had no consistent effects on TR and CS. Water stress has
77

78
been shown to affect growth (31,116), yield, WUE (10,11,40)
and physiological responses (25,126) in banana (Musa spp.
AAA). Relatively fewer studies have been reported on
temperature stress effects (63,142,143). An air temperature
of 33°C was reported as being optimum for growth and dry
weight partitioning in Cavendish banana (142) but the
investigators experienced some difficulty in separating the
stress effects of air temperature and induced vapor pressure
deficits. Investigations in banana involving both stress
factors simultaneously have not been reported. A similar
pattern of research exists for coffee, a major tropical crop
that belongs to the same subfamily as ixora (Ixora chinensis
L. ' Maui') (6).
This study investigated the effects of four RZTs and two
irrigation volumes on growth, physiology and carbohydrate
status of 'Grande Naine' banana and 'Maui' ixora under
greenhouse and growth room conditions.
Materials and Methods
Plant Materials and General Cultural Procedures
Ten- to 12-cm tissue-cultured 'Grande Naine' banana
plants and 12 to 15 cm uniform ixora rooted cuttings were
obtained from commercial nurseries. Plants were hardened in
intermittent mist (6 secs min-1) under 80% light exclusion
for one week and then moved to 40% light exclusion for
another week. Plants were transplanted to 6.5-cm diameter x
25.0-cm tall conical containers (555 cm3) using Metro-Mix 300

79
growth medium (W.R. Grace and Co., Cambridge, MA). Plants
were then moved to the experimental greenhouse or growth room
where they were watered to container capacity daily and
allowed to acclimatize for one week prior to the initiation
of the experiment.
Experiment 1--Greenhouse
Experiment 1 was initiated in August, 1985, in an air-
conditioned greenhouse in which maximum irradiance levels
ranged from 600 to 800 umol m"2 S-1 ppfd, and 25° to 30°C day
and 18° to 21°C night temperatures were maintained. Relative
humidity was not controlled and varied from 40% to 80%
(Figure 5-1).
Two irrigation volumes and four RZT treatments were
factorially combined in a split plot design experiment.
Plants were watered daily at 2200 hr with 50+5 ml (Wl) and
100+_10 ml (W2) per container ( 555 cm3). These volumes
represented 60% to 70% and 85% to 100% container capacity
(CC), respectively. Water was applied automatically through
drip tubes by a battery-operated controller (Water Watch
Corp. Seattle, WA). The Wl and W2 irrigation treatments were
applied by one and two 6-cm dramm irrigation rings/container,
respectively.
RZTs of 28+_l°, 33 +_10, 38+_l° and 43+_l°C were established
in air-bath boxes as described in Chapter IV. Each
temperature box contained six ixora and six banana plants to
which the irrigation treatments Wl and W2 were randomly

PPFD lumol
Figure 5-1. Photosynthetic photon flux density and relative humidity in the
greenhouse during measurements of physiological parameters.
PERCENT RELRTIUE HUniDITY

81
assigned to three plants per species per treatment.
Temperature boxes were replicated three times. RZT treatments
were applied daily from 1000 to 1600 hr EDT and irrigation
was supplied at 2200 hr so that the imposition of temperature
and irrigation treatments did not overlap. Plants were
fertilized weekly with the appropriate volume of water and
concentrations of 20N-8.8K-16.6P fertilizer (Peters 20-20-20,
W.R. Grace and Co., Cambridge, MA) to apply 250 mg
N/container. Fertilizer was applied with a hand operated
injector (Chem-trol Inc, Kansas City, KS).
Plant height, leaf area, leaf number and stem diameter
for banana, and plant height, plant width, shoot number and
total axillary shoot length for ixora were recorded weekly.
Plant height was considered the distance from the soil
surface to the leaf apex in banana and to the stem tip in
ixora. Banana stem diameter was measured at pot rim level and
leaf area of the third newest leaf was calculated from 0.65
of leaf length multiplied by leaf width (127). Leaf numbers
were derived from the number of whole green functional leaves
and emerging leaves were assigned a value of 0.25, 0.50 or
0.75 depending on the fraction of the leaf lamina fully
exposed.
Ten weeks after the initiation of the experiment, leaf
photosynthesis (PS), leaf conductance (CS) and transpiration
(TR) were recorded on a cloudless day from 1000 to 1600 hr at
2-hr intervals using a portable photosynthesis system (Model
LI-6000, LI-C0R Inc., Lincoln, NE). A 1-liter leaf cuvette

82
was used and the mean of eight 30-sec observations
constituted a measurement. A zero check of the analyzer was
performed between treatments within each replicate.
Simultaneous measurements of LWP were made using a pressure
chamber (PMS Instrument Model 600 Santa Barbara, CA) as
described by Barrett and Nell (9). The third most recently
expanded leaf in banana was selected for gas exchange
processes and LWP measurements. This leaf has been shown to
be the most responsive and the youngest with fully developed
stomata (127). The most recently-matured ixora leaves were
used for PS, CS and TR measurements but 4- to 5- cm shoot
tips were sampled for LWP.
After physiological measurements, leaf and root samples
were immediately collected for carbohydrate analysis. The
third newest leaf in banana and six of the most fully
developed leaves in ixora were sampled. A 5- to 7-g composite
sample of carefully washed roots was used for root
carbohydrate analysis. Plants were then separated into
leaves, stems and roots before drying for at least 48 hr in a
forced-air oven at 70°C prior to recording dry weights.
Ethanol-soluble sugars and starch were determined
according to the procedure used by Stamps (133) in which free
sugars were extracted from a 50-mg sample with 80% ethanol.
Leaf and root tissues were first freeze-dried at -40°C
(Freezemobi1e II, Virtis Co. Gardner, NY) for at least 72 hr
and stored in a desiccator. Tissue was then ground through a
20-mesh screen (Wiley Mill, Philadelphia, PA) and a 50-mg

83
sample was boiled in 8 ml of 80% ethanol and centrifuged for
1 hr at 10,500 rpm (Model HT Centrifuge, International
Equipment Co., Division of Damon). A 0.5-ml sample of
supernatant was diluted from 10 to 20 times with deionized
water. One half milliliter of 5% phenol and 2.5 ml
concentrated H2SO4 were added to 0.5 ml of each diluted
sample and the mixture was allowed to cool. Absorbance was
then measured at 490 nm using a spectrophotometer. Free sugar
concentrations were determined from plots of absorbance by
standard solutions of D-glucose and deionized water. Starch
content was determined by incubating the centrifuged pellet
at 34°C for 12 hr in 5 ml of an enzyme solution containing
sodium phosphate mono- and dibasic, alpha-amylase,
amyloglucosidase and anhydrous calcium chloride. After
incubation, samples were centrifuged for 1 hr and absorbance
determined using the same procedure as above.
The experiment was arranged in a split plot design with
three blocks in which RZT were main plots and irrigation
treatments represented subplots. Each species was treated and
analyzed as a separate experiment. Mean values of
physiological measurements were plotted across time to reveal
diurnal patterns and regression analyses of midday
physiological parameters over RZT were performed for each
irrigation volume treatment. Weekly leaf area data of banana
were used for growth over time plots and final growth and
carbohydrate measurements were used for regression analyses
over RZT and irrigation treatments. Single degree-of-freedom

84
orthogonal comparisons between treatment means were performed
and partitioning of interactions for linear, quadratic and
cubic models for final growth and carbohydrate data were made
using the General Linear Model procedure of the Statistical
Analysis System (134).
Experiment 2--Growth Room
In order to further monitor the physiological responses
of banana and ixora to RZT and irrigation treatments under
more precisely controlled environmental conditions,
experiment 1 was repeated in a walk-in growth room in August
and September, 1986. The growth room was described in
Chapters III and IV. Irrigation and RZT treatments were the
same as for experiment 1 but root heat tube (RHT) containers
as described in Chapter III were used. Similar plant
materials and cultural procedures were used as for experiment
1 except plants were first transplanted for one week into 4-
mi1 7.5 cm diameter x 22.5 cm tall clear plastic bags (1200
cm3) that were inserted into similar sized lengths of PVC
tubes. One week prior to the initiation of treatments, the
plastic bags were transferred to RHTs in the growth room.
Irrigation volumes of 75jj8 ml (Wl) and 150 +_15 ml (W2) per
container were applied daily at 2200 hr as in experiment 1.
These volumes provided 60% to 70% container capacity (CC) and
85% to 100% CC, respectively. Water was applied automatically
by drip irrigation and one and two 1-mm drip tubes per
container constituted the Wl and W2 treatments, respectively.

85
There were 16 RHTs per electronic controller, four of
which were randomly assigned to each RZT of 28 + 0.3°, 33+_0.3°,
38 + 0.3° and 43+_0.3°C. There were six replicate plants per
RZT-irrigation treatment arranged in a randomized complete
block design with three blocks. Species were organized and
treated as separate experiments. RZT treatments were applied
from 1000 hr to 1600 hr daily.
Six weeks later PS, CS, TR and LWP were measured at 2 hr
intervals from 0800 to 1800 hr and growth data and samples
for carbohydrate analysis were taken as described in the
greenhouse experiment. Chlorophyll concentration of leaves
used for physiological measurements was recorded using a Spad
501 chlorophyll light absorbance meter (Minolta Corp. Ramsey,
NJ) and expressed as umol m“2.
Means and standard errors were calculated from
physiological measurements of six replicate plants at each
sampling time and are shown with data points. Single degree-
of-freedom orthogonal comparisons between treatment means and
partitioning of interactions for linear, quadratic and cubic
models for growth, physiological and carbohydrate data were
made using the General Linear Model procedure of the
Statistical Analysis System (134).

86
Results and Discussion
Experiment l--Greenhouse
Physiological responses--banana. Under the W1 irrigation
volume, maximum PS at 1200 and 1400 hrs was 0.285 and 0.263
mg CO2 m“2 s"1, respectively (Figure 5-2A). Diurnal patterns
for CS and TR appeared to parallel each other under both
irrigation levels with maximum levels for both parameters
occurring at 1200 and 1400 hr (Figure 5-2, 5-3). LWPs were
generally higher in plants at the 28° and 33°C RZTs under the
W1 irrigation volume except at midday when LWP averaged
-0.71 MPa over all RZTs (Figure 5-2D).
At the higher irrigation level, W2, a maximum PS of
0.330 mg CO2 m-2 s“was measured for the 33°C RZT treated
plants at midday (Figure 5-3A) and another similar peak of
0.301 mg CO2 m-2 s“* occurred in the 38°C RZT treated plants
at 1400 hr. PS was significantly reduced at midday and
thereafter by the 43°C RZT. LWPs were generally higher in the
33° and 38°C RZT treated plants compared to those observed at
28° and 43°C between 1000 and 1400 hr (Figure 5-3D).
Regression analyses of midday PS over RZT and irrigation
treatments revealed significant effects of RZT but not
irrigation treatments (Figure 5-4A). PS was highest at the
28° and 33°C RZT treatments with progressive reductions at
the 38° and 43°C RZTs. While highest midday PS was noted for
the 28° and 33°C RZTs, diurnal patterns showed a lag in
response for the 38°C RZT with peak PS occurring after midday
(Figure 5-2A, 5-3A). Since there were also corresponding

Figure 5-2. Effects of four root-zone temperatures and a 50+5 ml daily irrigation
volume per 555 cm3 container on the diurnal physiological responses of 'Grande
Naine* banana grown under greenhouse conditions. A. leaf photosynthesis B. leaf
conductance C. transpiration and D. leaf water potential. Points are the means
of six replicate plants and vertical bars represent the SE.

■ M I.—.I II I I . I I I .-III. I ■ I ■ A ■ “1.0 I
1000 1200 1400 1600 1000 1200
TIME (hrs) TIME (hrs)
o
TRANSPRAT10N (mg H20 m-2 9-X)
4k ® M CD
PHOTOSYNTHESIS (mg 002 m-2 s-1)
o 3 & fc k
T
LEAF WATER POTENTIAL (MPa)
¿ ¿ ¿
b b ^
o
PO
LEAF CONDUCTANCE (cm s-1)
i S £ 3
88

Figure 5-3. Effects of four root-zone temperatures and a 100+_10 ml daily
irrigation volume per 555 cm^ container on the diurnal physiological responses of
'Grande Naine' banana grown under greenhouse conditions. A. leaf photosynthesis
B. leaf conductance C.transpirati on and D. leaf water potential. Points are the
means of six replicate plants and vertical bars represent the SE.

TRANSPIRATION (mg H20 m-2 8-1)
o 4k ao w oí 8
PHOTOSYNTHESIS (mg 002 m-2 8-1)
o
o
o
ZÍ-L
m8
CD
â– P
-r
k
k
6
ro
o
ro
5 8
b
ca
T"
O
-r-
ro
03
06

Figure 5-4. Regressions of midday physiological responses of container-grown
'Grande Naine' banana over four root-zone temperatures and two irrigation volumes
under greenhouse conditions. Main effects of root-zone temperature on leaf
photosynthesis are presented in A and interactive effects of root-zone
temperature and irrigation volume on leaf conductance, transpiration and leaf
water potential are presented in B, C and D, respectively. W1 (O) = 50 + 5 ml , W2
(â–¡) = 100+10 ml daily per 555 cm^ container.

ROOT-ZONE TEMPERATURE (°C> ROOT-ZONE TEMPERATURE (°C)
TRANSPIRATION (mg H20
-2 ,-ll
-2 ,-lj
Q
LEAF PHOTOSYNTHESIS (mg C02 m
rO
O
+
u
Q
+
n
>
Z6

93
shifts in CS and TR, there might have been a RZT-induced
delay in stomatal opening in the morning hours.
There were significant interactions between irrigation
treatments and RZT on midday CS, TR and LWP (Figure 5-4B, 5-
4C, 5-4D). For plants under the W1 irrigation level, a
quadratic trend was observed for CS and TR. CS and TR were
generally increased by the 33°C RZT compared to the other
RZTs. At the W2 irrigation level, TR and CS decreased
linearly with increasing RZT.
Irrigation treatments altered the influence of RZT on
midday LWP with the W2 level resulting in higher LWPs at all
RZTs except in plants exposed to the 28°C treatment (Figure
5-4D). RZTs above 28°C, increased plant water status under
well-watered conditions but a correspondí'ng increase in gas
exchange processes did not occur. This suggested a direct
effect on stomatal opening by high RZTs which was not
influenced by LWP.
Growth responses--banana. Plant height, stem diameter,
leaf length and leaf number did not differ with RZT or
irrigation treatments but width and area of the third newest
leaf were significantly reduced by RZT (Table 5-1). Effects
of the 43°C RZT became evident as early as two weeks after
initiation of treatments (Figure 5-5) with induced reductions
in leaf area expansion by the 38° and 43°C RZTs occurring

94
Table 5-1. Growth components of 'Grande Naine' banana measured
after 10 weeks at four root-zone temperatures and two
irrigation volumes under greenhouse conditions.
Stress
treatments
Plant
height
(cm)
No. of
leaves
Stem
d i a m.
(cm)
Leaf2
length
(cm)
Leaf2
width
(cm)
Leaf2
area
(cm2)
RZT (°C)
28
44.0
11.6
2.6
23.7
10.1
155.9
33
43.6
11.5
2.4
22.8
10.5
155.7
38
43.2
11.9
2.6
23.3
09.8
148.4
43
43.7
12.2
2.6
21.2
09.2
126.7
Irrigation
volume (IRV)
wiy
42.6
11.6
2.6
22.6
09.9
145.4
W 2
44.2
11.8
2.5
22.9
09.8
145.9
Significancex
RZT
NS
NS
NS
NS
Q*
Q*
IRV
NS
NS
NS
NS
NS
NS
Measurements
taken on
the third
newest
leaf.
m - 50+5 ml, W 2 - 100+^10 ml per container daily.
Statistical differences between treatments resulting from
orthogonal comparisons. Best fit models were Quadratic (Q).
*, NS - significant at the 5% probability level and
nonsignificant, respectively .

Figure 5-5. Effects of four root-zone temperatures across two irrigation volumes
(5 0+_5 ml and 100+_10 ml daily per 555 cm^ container) on leaf area of the third
newest leaf in container-grown 'Grande Naine' banana measured over 10 weeks under
greenhouse conditions. Points are the means of six replicate plants and vertical
bars represent the SE.

LEAF AREA (cm2
160
140
120
100
80
60
5 6 7
TIME (UEEKS)
8 8 10
cr>

97
throughout the experiment. At 10 weeks the effects of the
high RZTs on leaf area were being moderated since many roots
were growing at or above the soil level of the containers and
escaping RZT treatments. Regression analysis of final leaf
area with RZT indicated a quadratic relationship
(y=-52.26+13.497x-0.217x2; r ^ = o. 9 7 ) with reductions in leaf
area at the 38° and 43°C RZTs without irrigation effects
(Table 5-1). Reductions in leaf width was the major
contributor to reductions in leaf area.
Shoot, stem and total plant dry weights were not
influenced by RZT or irrigation treatments (Table 5-2).
However, effects of RZT on root dry weight and thus
shoot/root ratio were altered by irrigation treatments
(Figure 5-6A, 5-6B). The general RZT-induced increase in root
dry weight of the greenhouse-grown banana plants appeared to
contradict the decrease in root weight expected under similar
RZT treatments (30,39,44,65). By the end of the 10-week
experiment, plants were somewhat pot-bound and roots of
plants exposed to the higher RZTs were located mainly in the
upper portion of the container. Due to the gradient of +1.0°C
within containers, as noted previously, these roots may have
been exposed to the lower limits of each RZT treatment.
Regression analyses indicated that root dry weight was
increased by the 38° and 43°C RZTs under the W1 treatment but
only by the 38°C RZT in the W2 treated plants (Figure 5-6B).
The 43°C RZT in the W2 treatments could have disrupted the
absorptive capacity of banana roots thus creating a flooding-

98
Table 5-2. Dry weight components of 'Grande Naine' banana
measured after 10 weeks at four root-zone temperatures and two
irrigation volumes under greenhouse conditions.
Stress
treatments
Stem
dry wt
(g)
Shoot
dry wt
(g)
Total plant
dry wt
(g)
RZT (oc)
28
14.4
20.1
25.8
33
12.9
17.9
23.7
38
14.2
20.3
29.3
43
12.9
18.5
25.9
Irrigation
volume (IRV)
W lz
13.7
19.3
26.5
W 2
13.6
19.1
24.9
Significancey
RZT
NS
NS
NS
IRV
NS
NS
NS
ZW1 - 50+5 ml
, W2 - 100+10 ml per
container
daily.
ystatistical differences between
orthogonal comparisons.
treatments
resulting from
NS - nonsignificant at the 5% probability level.

99
u2 y =-3)0.14+27.91x-0.816x2+0. 00?8Sx3
r2.0. 94
-I 1 h
28 33 38
ROOT-ZONE TEMPERATURE (°C)
43
ROOT-ZONE TEnPERflTURE l0C)
Figure 5-6. Effects of four root-zone temperatures and two
irrigation volumes on A. shoot/root ratio and B. root dry
weight of container-grown 'Grande Naine' banana under
greenhouse conditions. W1 (O) = 50+_5 ml, W2 (â–¡) = 100+10 ml
per 555 cm^ container daily. Points are irrigation treatment
means.

100
like situation under the high irrigation treatment and
resulting in restricted root growth. A similar condition was
reported for well-watered container-grown eggplants (Solanum
me 1 ongena L.) grown at 40°C RZT which resulted in decreased
root dry weight (118).
Root morphology was also influenced. Roots were lacking
distinct tips and were brown and less succulent in plants at
the 38° and 43°C RZTs than at the lower RZTs. Suberization of
roots at high RZT (22° to 350C) was observed in ryegrass (Poa
pratensis L.) (32), peach (Prun us pérsica Batch.) (99), and
rose (Rosa sp. L.) (124).
Carbohydrate analysis--banana. Absolute shoot
carbohydrate content was not influenced by stress treatments
under greenhouse conditions but there were significant RZT
effects on root sugar and shoot sugar/root sugar ratios
(Table 5-3). RZT and irrigation volumes also affected root
sugar/starch ratio (Figure 5-7). Root sugar content was
greatest in plants grown at the 33° and 38°C RZTs. Shoot
sugar/root sugar ratios revealed a corresponding decrease by
the 33° and 38°C RZT (Table 5-3) and plants treated with
these RZTs generally had better water status as reflected by
LWP (Figure 5-4D). Increased shoot sugar or shoot sugar/root
sugar ratio have been implicated in the process of osmotic
adjustment (57,74,105). Sugars not needed for maintaining
turgor in plants at the 33° and 38°C RZTs could be available
for translocation to the roots thus giving higher root sugar

101
Table 5-3. Shoot and root carbohydrate distribution in 'Grande
Naine' banana measured after 10 weeks at four root-zone
temperatures and two irrigation volumes under greenhouse
conditions.
Stress
treatments
Shoot
sugar
(% dwt)
Shoot
starch
(% dwt)
Root
sugar
{% dwt)
Root
starch
{% dwt)
SSUG2
mnr
RZT (oc)
28
3.5
3.8
2.0
3.9
2.0
33
3.1
3.4
3.3
3.2
1.0
38
3.0
3.5
3.4
3.5
0.9
43
3.6
3.3
1.5
3.0
2.5
Irrigation
volume (IRV)
wiy
3.3
3.7
2.4
3.7
1.6
W 2
3.3
3.3
2.6
3.1
1.5
Significancex
RZT
NS
NS
Q**
NS
Q**
IRV
NS
NS
NS
NS
NS
ZSSUG - shoot
m - 50+5 ml
sugar, RSUG - root
, W2 - 100+10 ml per
sugar.
container
daily.
xStatistical
differences
between
treatments
resulting
from
orthogonal comparisons.
Best fit
models were Quadratic (Q).
**, NS - significant at the 1% probability level and
nonsignificant, respectively.

Figure 5-7. Effects of four root-zone temperatures and two irrigation volumes on
root sugar/starch content of 'Grande Naine' banana grown under greenhouse
conditions. W1 (O) = 50+_5 ml, W2 (â–¡) = 100+_10 ml daily per 555 cm3 container.
Points are irrigation treatment means.

ROOT-ZONE TEMPERATURE (°C)
ROOT SUGAR/STARCH RATIO
eoi
+0.727x-

104
concentrations at these RZTs. Partitioning between sugar and
starch in the roots was influenced by irrigation levels with
a higher root sugar/starch ratio in the W2 treated plants
(Figure 5-7). This agrees with other reports in which
increased starch to sugar conversion in the roots occurred
under well-watered conditions (106).
Physiological responses--ixora. PS of plants receiving
the W1 irrigation volume was generally higher at 1400 and
1600 hr if grown at 33° and 38°C RZTs compared to 28° and
43°C (Figure 5-8A). Maximum CS and TR occurred at midday for
the 28°C RZT treated plants and at 1400 hr for plants at the
330 and 38oc RZTs (FigUre 5-8B, 5-8C). The 43°C RZT reduced
CS and TR at midday and CS at 1400 hr. LWP declined from 1000
hr to midday at the lower irrigation level in plants at the
28°, 33° and 38°C RZTs, but increased to early morning levels
by 1600 hr (Figure 5-8D). Midday LWP at 43°C RZT was
apparently influenced by midday decreases in CS and TR and
did not decline to as low values as the other treatments at
midday.
Diurnal patterns of measured parameters in plants under
the higher irrigation volume were similar to those under the
W1 irrigation treatment. Maximum PS occurred at 1400 hr in
the 33°C RZT treated plants and in the 33° and 38°C RZT
treated plants at 1600 hr (Figure 5-9A). CS and TR were
significantly higher between 1200 and 1400 hr for the 28° and
33°C treated plants compared with the other RZTs

Figure 5-8. Effects of four root-zone temperatures and a 50+_5 ml daily irrigation
volume per 555 cm^ container on the diurnal physiological responses of Ixora
chinensis L. 'Maui' grown under greenhouse conditions. A. leaf photosynthesis B.
leaf conductance C. transpiration and D. leaf water potential. Points are the
means of six replicate plants and vertical bars represent the SE.

1000 1200 1400 1600 1000 1200 1400
TIME (hrs) TIME (hrs)
TRANSPfiATlON (mg H20 m-2 s-1)
^ ^ ® m ® 8
i
Kj
i
b
¿
¿
PHOTOSYNTHESIS (mg C02 m-2 s-1)
b b *-* i* K?
O 5 09 n> as O
OJ
901

Figure 5-9. Effects of four root-zone temperatures and a 100+10 ml daily
irrigation volume per 555 cm3 container on the diurnal physiological responses
Ixora chinensis L. 'Maui' grown under greenhouse conditions. A. leaf
photosynthesis B. leaf conductance C. transpiration and D. leaf water
potential. Points are the means of six replicate plants and vertical bars
represent the SE.

TRANSPRATON (mg H20 m-2 s-1)
„ ^ K)
O W IO 05 O
T
O
O
o
fO
á §
m
CO _1
A .
8
T"
T
i-
PH OTO SYNTHESIS (mg C02 m-2 s-1)
o 6 8 * ® §
ro
T"
05
-I—
^ co co ro
co oo co a>
o o o o
O O O O
LEAF WATER POTENTIAL (MPa)
A ¿ ¿ ¿
b bo ® k>
o
o
o
S.ro
m§
09
â– U
8
LEAF CONDUCTANCE (cm s-1)
b
GO
ro
80T

109
(Figure 5-9 B, 5-9C). Decreased LWP corresponded to increases
in TR and CS at midday and LWP increased as TR and CS
declined at 1600 hr (Figure 5-9D). Plants at the 43°C RZT had
lowest LWPs at 1000 and 1200 hr and the 38°C RZT treated
plants had highest LWP at midday.
Significant effects of RZT on PS, CS and TR were
observed when midday values of these parameters were
regressed over RZT but irrigation treatments did not alter
these effects (Figure 5-10). Quadratic trends occurred for PS
and CS with significant decreases at the 43°C RZT (Figure 5-
10A, 5-10B) and TR decreased linearly with increasing RZT
(Figure 5-10C). Regression analysis for LWP indicated
significant interactions of RZT and irrigation treatments
(Figure 5-10D). Midday LWP of plants receiving the W2
irrigation treatment were significantly higher than those at
the W1 level except at the 43°C RZT (Figure 5-10D). The 43°C
RZT,therefore, decreased all measured gas exchange processes
at midday in ixora and increased water volume did not improve
plant water status. Since the 43°C RZT also appeared
supraoptimal after two weeks of RZT stress treatments
(Chapter IV), there apparently was little conditioning in
ixora to this high RZT.
Growth responses--ixora. Except for axillary shoot
number, most growth components and the number of flower buds
in the greenhouse-grown plants were significantly affected by
RZT but not by irrigation treatments (Table 5-4). While

Figure 5-10. Regressions of midday physiological responses of container-grown
Ixora chinensis L. 'Maui' over four root-zone temperatures and two irrigation
volumes under greenhouse conditions. Main effects of root-zone temperature on
leaf photosynthesis, leaf conductance and transpiration are presented in A, B and
C, respectively and interactive effects of root-zone temperature and irrigation
volume on leaf water potential are presented in D. W1 (O ) = 50+5 ml , W2 (â–¡ ) =
100+10 ml daily per 555 cm^ container.

ROOT-ZONE TEMPERATURE (°CI ROOT-ZONE TEMPERATURE (°CI
ITT

112
Table 5-4. Growth components of Ixora chinensis L. 'Maui'
measured after 10 weeks at four root-zone temperatures and two
irrigation volumes under greenhouse conditions.
Stress
treatments
Plant
height
(cm)
Plant
width
(cm)
No. of
axillary
shoots
Axillary
shoot length
(cm)
No. of
f1ower
buds
RZT (oC)
28
32.5
19.4
2.8
37.3
1.2
33
40.8
24.0
3.2
52.9
0.6
38
43.0
25.2
3.3
61.5
0.2
43
37.1
22.8
2.6
51.2
0.2
Irrigation
volume (IRV)
W lz
39.1
22.5
2.8
50.4
0.6
W 2
37.6
23.2
3.1
51.1
0.5
SignificanceV
RZT
Q**
Q*
NS
Q**
Q**
IRV
NS
NS
NS
NS
NS
ZW1 - 50+5 ml, W2 - 100+_10 ml per container daily.
ystatistical differences between treatments resulting from
orthogonal comparisons. Best fit models were Quadratic (Q).
**, *, NS - significant at the 1%, and 5% probability levels
and nonsignificant, respectively.

113
growth parameters were increased (Figure 5-11A, 5-11B, 5-
11C), flower bud number declined at RZTs above 28°C (Figure
5-11D). Since flowering and subsequent axillary growth
occurred in the 28°C treated plants midway in the study,
axillary shoot length (Figure 5-11C) but not shoot number
(Table 5-4) was significantly different at 10 weeks.
There were interactive effects of RZT and irrigation
volume on shoot, total plant and root dry weights (Figure 5-
12A, 5-12B, 5-12D) but only RZT effects on shoot/root ratio
(Figure 5-12C). Shoot and total plant dry weights increased
at RZTs above 28°C under the W2 irrigation level but these
parameters were maximum at 38°C RZT under the W1 irrigation
treatment (Figure 5-12A, 5-12B). All dry weight parameters
were significantly increased by irrigation at the 33°C RZT.
While the higher irrigation level overcame the stress effects
of the 43°C RZT on shoot and total plant dry weight (Figure
5-12A, 5-12B), it did not increase root dry weight over that
in the W1 treated plants at 38° and 43°C RZT (Figure 5-12D).
Larger shoots with reduced roots at the higher RZTs
(Figure 5-12C) would imply a RZT-induced increase in root
absorption and/or increased assimilate supply. However,
trends in TR (Figure 5-10C) and PS (Figure 5-10A) suggested
that neither water absorption nor assimilate production was
increased in plants at RZTs above 33°C. This leaves the
possibility of a RZT-induced increase in hormonal synthesis
(64,76,91) and/or export from the roots (91,129) as a cause
for the increased axillary shoot growth. An increased

Figure 5-11. Main effects of four root-zone temperatures across two irrigation
volumes (50+5 ml and 100+_10 ml daily per 555 cm^ container) on growth of Ixora
chinensis L7 'Maui' grown under greenhouse conditions. A. plant height B. plant
width C. axillary shoot length and D. number of flower buds. Points are the means
of 12 replicate plants.

ROOT-ZONE TEMPERATURE I°C1 ROOT-ZONE TEMPERATURE l°C)
AXILLARY SHOOT LENGTH (cm)
PLANT HEIGHT (cm)
SIT

Figure 5-12. Effects of four root-zone temperatures and two irrigation volumes on
dry weight components of container-grown Ixora chinensis L. 'Maui' under
greenhouse conditions. Interactive effects of root-zone temperature and
irrigation volume on shoot dry weight, total plant dry weight and root dry weight
are presented in A, B and D, respectively and main effects of root-zone
temperature on shoot/root ratio are presented in C. W1 (O) = 50+5 ml, W2 (â–¡ ) =
100+_10 ml daily per 555 cm3 container. Points represent irrigation treatment
means in A, B and D and the means of 12 replicate plants in C.

ROOT-ZONE TEHPERRTURE (“Cl ROOT-ZONE TEflPERRTURE ("Cl
¿II

118
acropetal flow of root-synthesized cytokinin may be
postulated (21,129) since increased axillary shoot growth
(Figure 5-11C), apparently at the expense of flowering
(Figure 5-11D), was induced by increased RZT.
Carbohydrate analysis--ixora. Absolute shoot sugar and
shoot and root starch contents were not affected by RZT or
irrigation treatments under greenhouse conditions but root
sugar content decreased and thus shoot sugar/root sugar ratio
increased linearly with increasing RZT (Table 5-5). These
trends in carbohydrate status indicated a maintenance of
shoot sugar content with either decreased translocation to
the roots (96) or increased respiration in the roots (58)
with increasing RZT.
PS did not increase in plants at the 38°C RZT compared
to lower RZTs and declined at the 43°C RZT (Figure 5-10A).
However, demand for photosynthates in plants at the 38°C and
43°C RZTs would be increased by the increased sink strength
of RZT-induced axillary shoot growth. Under these conditions,
there would probably be less available soluble sugars for
translocation to the roots, hence the decreased root sugar
content. Increased root respiration at high RZT with less
available substrates could also have contributed to the
decreased root dry weight at the 43°C RZT (Figure 5-12B).

119
Table 5-5. Shoot and root carbohydrate distribution in Ixora
chinensis L. 'Maui' after 10 weeks at four root-zone
temperature s
conditions.
and two
irrigation
volume s
under greenhouse
Stress
treatments
Shoot
sugar
{% dwt)
Shoot
starch
(% dwt)
Root
sugar
(% dwt)
Root
starch
{% dwt)
Shoot sugar
root sugar
RZT (oc)
28
4.4
3.2
5.7
3.6
0.8
33
5.9
2.6
3.8
3.5
1.5
38
4.8
2.7
3.8
3.2
1.3
43
5.0
2.7
3.2
2.9
1.6
Irrigation
volume (IRV)
Wlz
5.2
2.6
4.1
3.1
1.4
W 2
4.9
3.1
4.2
3.5
1.2
SignificanceV
RZT
NS
NS
★ ★
NS
^ ★ ★
IRV
NS
NS
NS
NS
NS
ZW1 - 50+5 ml, W2 - 100+_10 ml per container daily.
ystatistical differences between treatments resulting from
orthogonal comparisons. Best fit models were Linear (L).
**, NS - significant at the 1% probability level and
non significant,respectively.

120
Experiment 2--Growth Room
Physiological responses--banana. Under the W1 irrigation
volume, plants at the 38°C RZT generally achieved higher
daily PS compared to the other RZTs with maximum PS rates of
0.485 and 0.470 mg C02 m-2 s_1 at 1200 and 1400 hr,
respectively (Figure 5-13A). Maximum PS recorded at two weeks
(Figure 4-3A, Chapter IV) in the growth room, also occurred
in 38°C RZT treated plants. High PS in plants exposed to the
38°C RZT for six weeks under reduced irrigation may suggest
that this RZT is within the optimum range for PS in banana.
The 43°C RZT resulted in comparatively lower PS at two and
six weeks after treatment imposition, indicating that there
was no evidence of conditioning to this extreme RZT
treatment.
Differences in CS and TR were not discernible between
RZTs except for the consistent reductions at midday and 1400
hr in the 43°C treatment. The 28°C RZT-treated plants had
significantly higher LWP at 0800 hr and midday.
Under the W2 irrigation level, the 33°C RZT treated
plants had relatively higher midday PS than those at other
RZTs (Figure 5-14A) and this was related to a correspondingly
higher TR at this RZT (Figure 5-14C). CS was significantly
reduced at midday and 1400 hr for plants under the 38° and
43°C RZT (Figure 5-14B) and there was a corresponding
decrease in TR for the 43°C RZT treated plants (5-14C).
These observations suggested an inhibitory effect on stomatal
opening by the 43°C RZT regardless of irrigation treatment

Figure 5-13. Effects of four root-zone temperatures and a 75+8 ml daily
irrigation volume per 1200 cm^ container on the diurnal physiological responses
of container-grown 'Grande Naine' banana under growth room conditions. A. leaf
photosynthesis B. leaf conductance C. transpiration and D. leaf water
potential. Points are the means of six replicate plants and vertical bars
represent the SE.

ZZT

Figure 5-14. Effects of four root-zone temperatures and a 150 +_15 ml daily
irrigation volume per 1200 cm^ container on the diurnal physiological responses
of container-grown 'Grande Naine' banana under growth room conditions. A. leaf
photosynthesis B. leaf conductance C. transpiration and D. leaf water
potential. Points are the means of six replicate plants and vertical bars
represent the SE.

LEAF
i
â– b
(o
WATER POTENTIAL (MPa)
¿
a>
¿>
ó>
LEAF CONDUCTANCE (cm s-1)
o % $ S &

125
and an inhibition by the 38°C RZT under the higher irrigation
level. RZTs above 28°C RZT reduced LWP during the morning
hours and increased irrigation did not appear to alter this
effect (Figure 5-13D, 5-14D).
Regression analyses of midday PS, CS and TR showed
significant effects of RZT and irrigation treatments (Figure
5-15). Maximum midday PS was recorded in plants at the 33°C
RZT under the W2 irrigation level with the 38° and 43°C RZT
treated plants having midday PS similar to that at 28°C. The
maximum midday PS in plants under the W1 irrigation was
recorded at 38°C RZT but this was 20% lower than PS in plants
at W2 and 33°C RZT (Figure 5-15A). RZT and irrigation
interactions also occurred for CS and TR with significant
increases under the W2 irrigation at the 28° and 33°C RZTs
(Figure 5-15B, 5-15C). RZTs above 28°C generally decreased
LWP, but increased irrigation increased LWP except at the
28°C RZT (Figure 5-15D). The positive effect of increased
irrigation on LWP at 38° and 43°C RZT but not on the gas
exchange processes is somewhat anomalous. As noted previously
there may have been some direct RZT effect on stomatal
opening at these RZTs independent of leaf water status.
Under growth room conditions, irrigation and RZT
interactions indicated that if growth medium temperature for
banana is maintained at 33°C, increased irrigation may be
beneficial. At 38°C, increased watering may increase plant
water status but not necessarily net carbon balance.

Figure 5-15. Regressions of midday physiological responses of container-grown
'Grande Naine' banana over four root-zone temperatures and two irrigation volumes
under growth room conditions. A. leaf photosynthesis B. leaf conductance C.
transpiration and D. leaf water potential. W1 (O ) = 75+8 ml , W2 (â–¡ ) = 150+JL5
ml daily per 1200 cm^ container. Points are irrigation treatment means.

ROOT-JONE TEMPERATURE (»C> R00T-30NE TEflEPRRTURE (°C>
LZI

128
Increased irrigation, however, moderates the effect of the
43°C RZT on gas exchange processes.
Growth responses--banana. Plant height, leaf number and
leaf length were not affected by RZT or irrigation treatments
but stem diameter was increased and leaf width and area were
decreased by increasing RZT under growth room conditions
(Table 5-6). The effects of RZT on leaf width and area were
generally similar to those that occurred in the greenhouse
study, although these parameters were linearly reduced by RZT
here as compared to the curvilinear relationship in the
greenhouse plants.
Dry weights were not altered by RZT or irrigation
volumes except for root dry weight and thus shoot/root ratio
which were affected by stress treatments with no interactive
treatment effects (Table 5-7). Root dry weight was reduced
linearly by increasing RZT with the lower irrigation volume
causing additional reduction. These results are in general
agreement with findings of high RZT on root growth in several
plants (30,39,44,65). The greater volume of the RHTs used in
this study compared to the containers used in the greenhouse
experiment was apparently more adequate for studying RZT
stress effects in banana plants. Root tips of plants at the
higher RZTs were absent and roots appeared more fibrous and
were less succulent at the higher RZTs than at 28° and 33°C
RZT. Stunting and browning effects on roots by high RZT has
been observed in rye grass (32), peach (99), and rose (124).

129
Table 5-6. Growth components of 'Grande Naine' banana measured
after six weeks at four root-zone temperatures and two
irrigation volumes under growth room conditions.
Stress
treatments
Plant
height
(cm)
No. of
leaves
Stem
d i a m.
(cm)
Leaf2
length
(cm)
Leaf2
width
(cm)
Leaf2
area
(cm2)
RZT (oc)
28
37.7
10.0
2.2
21.3
9.9
137.4
33
39.1
10.2
2.5
20.7
9.9
133.5
38
36.5
9.9
2.8
20.9
8.4
114.5
43
36.2
9.8
3.2
20.7
8.2
110.5
Irrigation
volume (IRV)
W1Y
36.7
9.8
2.6
21.1
9.3
127.4
W 2
37.7
10.2
2.7
20.8
9.0
122.0
S i g n i f i c a n c e x
RZT
NS
NS
★
NS
★
IRV
NS
NS
NS
NS
NS
NS
Measurements
taken on
the third
newest
leaf.
VWl - 75+8 ml, W2 - 150^15 ml per container daily.
Statistical differences between treatments resulting from
orthogonal comparisons. Best fit models were Linear (L).
**, NS - significant at the 1% probability level and
nonsignificant, respectively.

130
Table 5-7. Dry weight components of 'Grande Naine' banana
measured after six weeks at four root-zone temperatures and two
irrigation volumes under growth room conditions.
Stress
treatments
Stem
dry wt
(g)
Shoot
dry wt
(g)
Root
dry wt
(g)
Shoot/root Total
ratio plant dry wt
(g)
RZT (°c)
28
10.5
17.9
7.6
2.4
25.6
33
10.2
19.7
6.4
3.1
26.1
38
10.7
18.7
5.3
3.5
24.0
43
10.2
19.9
4.1
5.0
24.0
Irrigation
volume (IRV)
W lz
10.1
18.4
5.0
3.9
23.6
W 2
10.9
19.4
6.8
3.0
26.0
Significancey
RZT
NS
NS
^ ★
★ ★
NS
IRV
NS
NS
★ ★
NS
NS
ZW1 - 75+8 ml
, W2 - 150+15 ml per
container daily.
ystatistical
differences
between
treatments resulting
from
orthogonal comparisons.
Best fit
models
were Linear
(L).
**, NS - significant at the 1% probability level and
nonsignificant, respectively.

131
As in the greenhouse experiment (Table 5-1, Figure 5-5),
leaf area was also decreased by the higher RZTs in this study
and the trend in reduction paralleled that of leaf width
(Table 5-6). There were also no RZT effects on plant height,
which is basically a measure of total leaf length in the
banana plant. Anatomical studies revealed that the unfolding
banana leaf increases in width mainly through expansion of
groups of meristematic cells still evident within the leaf
lamina (130). These observations suggested a RZT-induced
effect on cell expansion and/or division rather than on cell
elongation.
Inhibition of leaf expansion by the 38° and 43°C RZT
treatments could have involved physiological mechanisms not
entirely associated with leaf water deficits. De Langhe et
a 1. (34), working on endogenous hormonal patterns in
developing banana plants, demonstrated that the root tips
were a major source of cytokinins in banana plants. Reduced
leaf expansion simultaneous with the loss of root tips
observed in the high RZT-treated plants under both
environmental conditions may imply a hormonal role in leaf
expansion. Since leaf width and not leaf length was affected
by high RZT, it may be hypothesized that root-synthesized
cytokinin was limiting in plants subjected to the 38° and
43°C RZTs. This could be validated in future studies by the
application of exogenous cytokinin to high RZT-treated plants
and monitoring the effects on leaf expansion.

132
Leaf chlorophyll content as measured by light absorbance
increased linearly with increasing RZT but was not affected
by irrigation treatment (Figure 5-16). Increased chlorophyll
content however, did not relate to increased PS (Figure 5-
15A), thus chlorophyll density may not have increased. Leaf
area was affected but dry weight was not altered by RZT.
These observations may suggest that a smaller but thicker
leaf was produced in response to high RZT.
Carbohydrate analysis--banana. Under the growth room
environment, root starch content decreased with increasing
RZT (Table 5-8) and there were interactive effects of RZT and
irrigation treatments on root sugar and root sugar/starch
ratio (Figure 5-17). Regression analysis indicated that root
sugar concentration decreased linearly with RZT at the W1
irrigation level but was cur vi 1inear 1y related to RZT at W2
(Figure 5-17A). RZTs above 28°C generally decreased LWP
(Figure 5-15D) thus possibly increasing the demand for
soluble sugars in the shoots for osmotic adjustment and
leaving less for translocation to the roots. Higher
irrigation moderated the RZT stress effects at 43°C (Figure
5-15) and roots were apparently not damaged. A high sink
demand of roots under this RZT and the W2 level could have
caused the noted root sugar content increase compared to the
330 an(j 380c RZTs even under comparable LWPs.

60
10
28 33 38 43
ROOT-ZONE TEHPERRTURE (°C)
Fiqure 5-16. Effects of four root-zone temperatures across two irrigation volumes
(75+8 ml and 150+15 ml daily per 1200 cm3 container) on chlorophyll concentration
of container-grown 'Grande Naine' banana under growth room conditions. Points are
the means of 12 replicate plants.
133

134
Table 5-8. Shoot and root carbohydrate distribution in 'Grande
Naine' banana measured after six weeks at four root-zone
temperatures and two irrigation volumes under growth room
conditions.
Stress
treatments
Shoot
sugar
(% dwt)
Shoot
starch
(% dwt)
Root
starch
(% dwt)
Shoot sugar
root sugar
RZT (oc)
28
4.9
4.9
8.3
0.8
33
4.0
4.4
8.0
0.7
38
4.0
4.6
7.7
0.9
43
3.7
4.0
6.8
1.1
Irrigation
volume (IRV)
Wlz
3.9
4.3
7.6
1.0
W 2
4.3
4.7
7.8
0.8
SignificanceY
RZT
NS
NS
★
NS
I R V
NS
NS
NS
NS
ZW1 - 75+8 ml, W2 - 15 0 +_15 ml per container daily.
ystatistical differences between treatments resulting from
orthogonal comparisons. Best fit models were Linear (L).
**, NS - significant at the 1% probability level and
nonsignificant, respectively.

Figure 5-17. Interactive effects of four root-zone temperatures and two
irrigation volumes on carbohydrate content of container-grown 'Grande Naine'
banana under growth room conditions. A. percent root sugar and B. root
sugar/starch ratio. W1 (O) = 75+8 ml, W2 (â–¡) = 150^15 ml daily per 1200 cm
container. Points are irrigation treatment means.

N)
O
PERCENT ROOT SUGAR
CD
O
o
o
+
+
9£I

137
Partitioning trends between sugar and starch in the
roots were affected by irrigation treatments (Figure 5-17B).
Well-watered conditions have been implicated in the
interconversion of starch to sugar in roots (107). The
expected increase in sink strength at higher RZT and the
effect of irrigation may have been responsible for the
quadratic trend in root sugar/starch under the W2 irrigation
treatment and the linear decrease in the W1 treated plants.
Starch content was linearly reduced with increasing RZT
irrespective of irrigation treatment (Table 5-8), reflecting
possible root starvation effects and the RZT-induced decrease
in root dry weight (Table 5-7).
Carbohydrate status and partitioning in banana appeared
to be related with RZT-induced changes in plant water status.
Turner and Lahav (142) reported significant shifts in
assimilate partitioning in banana plants subjected to
increasing air temperatures under growth room conditions. In
their study, however, neither LWP nor carbohydrate content
were measured and their results were based entirely on dry
weight determinations. Relating LWP with plant carbohydrate
status is probably a more direct method of explaining RZT-
induced shifts in partitioning patterns.
Physiological responses--ixora. PS rates were generally
higher in the growth room than in the greenhouse. However,
diurnal PS patterns were not greatly affected by RZT under
the W1 irrigation level in the growth room, except for the

138
reduced midday rates for the 43°C RZT treated plants (Figure
5-18A). This general pattern was also evident for CS (Figure
5-18B) and TR except that midday TR was also reduced by the
38°C RZT (Figure 5-18C). LWPs appeared to be influenced by
RZT and its effects on CS and TR. Transpirational water loss
did not reduce LWP at 28°C RZT compared to that in plants
grown at 33° and 38°C RZTs (Figure 5-18D). The 43°C RZT
inhibited stomatal opening directly but apparently did not
reduce the absorptive capacity of the roots thus resulting in
LWP comparable to that in the 28°C RZT treated plants. Linder
the W2 irrigation, effects of RZT on diurnal fluctuations
were less apparent than in plants with the lower irrigation
volume (Figure 5-19), but the 43°C RZT resulted in reduced PS
and decreased CS and TR at 1000 hr through midday.
There were no interactive effects of RZT and irrigation
treatments on midday PS, CS and TR but quadratic
relationships with RZT were noted (Figure 5-20A, 5-20B, 5-
20C). RZTs of 28°, 33° and 38°C resulted in similar midday
PS, CS and TR but there were discernible reductions for
plants at 43°C. This pattern of RZT effects on gas exchange
parameters was also apparent in plants in the greenhouse
study (Figure 5-10). Regression curves were not presented for
responses to RZT in the two week growth room stress study
(Chapter IV), but midday values of PS, CS and TR indicated a
similar trend as occurred at six weeks. These results
suggested that there was little conditioning in ixora to the

Figure 5-18. Effects of four root-zone temperatures and a 75+8 ml daily
irrigation volume per 1200 cm^ container on the diurnal physTological responses
of Ixora chinenesis L. 'Maui' grown under growth room conditions. A. leaf
photosynthesis B. leaf conductance C. transpiration and D. leaf water
potential. Points are the means of six replicate plants and vertical bars
represent the SE.

0800 1000 1200 1400 1600 1800
'Ü
-04
,-0.6
S’0*8
I
5S-1X)
-12
0800 1000 1200 1400
TIME (hrs)
1600 1800
140

Figure 5-19. Effects of four root-zone temperatures and a 150+15 ml daily
irrigation volume per 1200 cm3 container on the diurnal physiological responses
of Ixora c h i n e n s iis L. 'Maui' grown under growth room conditions. A. leaf
photosynthesi s ET7 leaf conductance C. transpiration and D. leaf water
potential. Points are the means of six replicate plants and vertical bars
represent the SE.

TR ANSPffWnON (mg H20 m-2 s-1)^
é 8 8
" 1 » « f
8
i
o
leaf WATER POTENTIAL (MPa)
i§I—t
PHOTOSYNTHESIS (mg C02 m-2 8-1)
o & fe S fe
» I I I

Figure 5-20. Regressions of midday physiological responses of container-grown
Ixora chinensis L. 'Maui' over four root-zone temperatures and two irrigation
volumes under growth room conditions. Main effects of root-zone temperature and
irrigation volume on leaf photosynthesis, leaf conductance and transpiration are
presented in A, B and C, respectively and interactive effects of root-zone
temperature and irrigation volume on leaf water potential are presented in D. W1
(O) = 75 + 8 ml , W2 (â–¡) = 150+15 ml daily per 1200 cm^ container.

ROOT-ZONE TEMPERATURE (°C) ROOT-ZONE TEMPERATURE I°C1
TRANSPIRATION (mg H20 m"2 i"1)
LEAF PHOTOSYNTHESIS (mg C02
2 ,-‘l

145
inhibitory effects of the 43°C RZT on stomatal opening. The
highest RZT however, appeared to limit root absorptive
capacity as reflected by midday LWP, more in two-week old
plants (Figure 3-3D) than at six weeks (Figure 5-18D).
The W2 irrigation volume treatment resulted in higher
midday LWP for plants at the 33° and 38°C RZTs (Figure 5-
20D). The W1 treatment apparently was inadequate for
maintenance of good water status of plants at 33° and 38°C
RZTs. While irrigation treatments significantly affected LWP,
there were no apparent effects on the gas exchange processes.
This discrepancy may have been due to the fact that while
apical shoot sections were sampled for LWP measurements,
recently matured leaves in ixora were selected for monitoring
gas exchange data. In the 28°C RZT treated plants, there were
not as many rapidly growing shoots as in higher RZT treated
plants and this probably influenced LWP results at the 28°C
RZT treatment.
Growth responses--!’xora. Under growth room conditions,
plant height and width were not influenced by stress
treatments but axillary shoot number were greatest at the 33°
and 38°C RZTs (Table 5-9). There were interactive effects of
RZT and irrigation volume on axillary shoot length (Figure 5-
21A). Shoot dry weight and thus shoot/root ratio increased
curvi1inearly with increasing RZT (Table 5-10) and RZT and
irrigation interactive effects on total plant and root dry

146
Table 5-9. Growth components of Ixora chinensis L. 'Maui'
measured after six weeks at four root-zone temperatures and two
irrigation volumes under growth room conditions.
Stress
treatments
Plant
height
(cm)
Plant
width
(cm)
No. of
axillary
shoots
RZT (°C)
28
30.8
16.7
2.0
33
32.1
16.8
3.2
38
31.6
17.0
3.6
43
28.7
15.0
2.7
Irrigation
volume (IRV)
W lz
30.2
16.3
2.8
W 2
31.1
16.4
2.9
Significancey
RZT
NS
NS
Q**
IRV
NS
NS
NS
ZW1 - 75+8 ml,
W2 - 150+15 ml
per container
daily.
ystatistical differences between treatments resulting from
orthogonal comparisons. Best fit models were Quadratic (Q).
**, NS - significant at the 1% probability level and
nonsignificant, respectively.

147
Table 5-10. Shoot dry weight, shoot/root ratio and chlorophyll
concentration of Ixora chinensis L. 'Maui' measured after six
weeks at four root-zone temperatures and two irrigation volumes
under growth room conditions.
Stress
treatments
Shoot
dry wt
(9)
Shoot/root
rati o
Chiorophyl1
concn
(umol m“2)
RZT (°C)
28
3.5
2.1
39.7
33
5.3
2.1
37.7
38
5.6
2.6
37.4
43
5.7
4.1
34.4
Irrigation
volume (IRV)
W lz
4.7
2.9
37.2
W 2
5.3
2.6
37.6
Significancey
RZT
Q*
/
Q**
NS
IRV
NS
NS
NS
ZW1 - 75+8 ml,
W2 - 150 + 15
ml per container
daily.
^Statistical differences between treatments resulting from
orthogonal comparisons. Best fit models were Quadratic (Q).
**, *, NS - significant at the 1%, and 5i probability levels
and nonsignificant, respectively.
i

Figure 5-21. Effects of four root-zone temperatures and two irrigation volumes on
growth of container-grown Ixora chinensis L. 'Maui' under growth room conditions.
A. axillary shoot length Í3. total plant dry weight and C. root dry weight.
W1 (O) = 75 + 8 ml, W2 (â–¡ ) = 150^15 ml daily per 1200 cm3 container. Points are
irrigation treatment means.

ROOT-ZONE TEI1PERRTURE (°C)
RXILLRRY SHOOT LENGTH tcm)
6t?T
S0O*
z*Pl*l *Q-x¿t>0 '0Í+CC '962-

150
weights were noted (Figure 5-21B, 5-21C). Stress treatments,
however, did not influence relative chlorophyll
concentrations (Table 5-10).
Stimulated axillary shoot length at 33° and 38°C RZTs
was further augmented by increased irrigation (Figure 5-21A).
Total plant (Figure 5-21B) and root (Figure 5-21C) dry weight
parameters showed quadratic relationships with RZT, with the
higher irrigation volume increasing dry weights at the 28°
and 33°C RZTs. Apparently, the higher light intensity and/or
longer photoperiod in the growth room retarded flowering in
ixora compared to the greenhouse conditions. The lack of
flowering prolonged apical dominance, particularly in the
28°C RZT treated plants. This resulted in an increased shoot
number in plants exposed to RZTs above 28°C (Table 5-9).
The induction of axillary shoot growth at 33°C RZT has
been implicated in studies by Franco (39) in coffee and Yusof
(148) in avocado. Stimulation of branching and vegetative
growth at the expense of flower production was reported for
container-grown peppers (44) at RZTs above 36°C. Hormonal
modification theories were suggested but not validated.
Carbohydrate analysis--ixora. Absolute shoot
carbohydrate content was not altered by stress treatments but
there were main effects of RZT on root sugar content and
shoot sugar/root sugar ratio (Table 5-11). While root sugar
content was maximum at 33° and 38°C RZTs, shoot sugar/root
sugar was a minimum at these RZTs. These relationships

151
Table 5-11. Shoot and root carbohydrate distribution in Ixora
chinensis L. 'Maui' after six weeks at four root-zone
temperatures and two irrigation volumes under growth room
conditions.
Stress
treatments
Shoot
sugar
{% dwt)
Shoot
starch
(% dwt)
Root
sugar
{% dwt)
Root
starch
(% dwt)
Shoot sugar
root sugar
RZT (oC)
28
4.4
3.1
2.2
8.2
2.0
33
3.7
3.4
4.6
6.9
0.8
38
4.3
4.4
4.6
9.7
0.9
43
4.3
4.0
2.7
5.8
1.6
Irrigation
volume (IRV)
W lz
4.3
3.6
3.5
7.9
1.3
W 2
4.1
3.8
3.2
7.3
1.3
Sgnificancey
RZT
NS
NS
Q**
NS
o**
IRV
NS
NS
NS
NS
NS
ZW1 - 75+8 ml, W2 - 150 +_15 ml per container daily.
yStatistical differences between treatments resulting from
orthogonal comparisons. Best fit models were Quadratic (Q).
**, NS - significant at the 1% probability level and
nonsignificant, respectively.

152
indicated that although absolute shoot sugar content was
unaffected by RZT, there was a tendency for increased
translocation of sugars to the roots at the 33° and 38°C
RZTs.
PS was generally higher in this study as compared to the
greenhouse experiment, and although induced axillary shoot
growth created extra sink demand, there probably was
assimilate supply to satisfy the increase in shoot sink
demand and that of the roots. Plants in the growth room did
not flower and this may have reduced the demand for sugars in
the shoots. The similarity in trends of root dry weight
(Figure 5-21C) and root sugar content (Table 5-11) over RZT
indicated that the supply of assimilate to the root was
probably more than adequate for root growth at the 33°C and
38°C RZT. In vegetative rice (Oryza sativa L.), a 38°C RZT
increased the flow of into the roots but decreased root
dry weights were attributed to the reduced incorporation of
carbon into protein and cell wall (97). In ixora under growth
room conditions, increasing RZT to 38°C apparently caused
increased translocation of sugars to the roots and induced
root growth. Environmental conditions appeared to highly
influence the effects of RZT treatments on carbohydrate
partitioning patterns.

CHAPTER VI
SUMMARY AND IMPLICATIONS
The major objective of this project was to evaluate the
effects of increasing RZT and irrigation volume,
independently and in interactive studies, on container-grown
'Grande Naine' banana (Musa spp. AAA) and Ixora chinensis L.
'Maui.' A portable CO2 gas analyzer system was used
effectively to measure gas exchange processes simultaneously
and the pressure chamber technique gave good indications of
plant water status. The more controlled conditions of the
walk-in growth room and the system of electronically
controlled RHTs allowed for a more concise study of plant
responses to RZT treatments than greenhouse conditions and
air bath boxes. The use of tissue-cultured banana plants
indicated that micropropagation can be an effective tool for
investigating physiological responses to stress factors.
Experiments on the independent effects of irrigation
volume reported in Chapter III showed conclusively that all
physiological parameters in banana were decreased by
decreasing irrigation volumes below the W3 (40+8 ml per 150
cm3 container daily; 85 to 100% CC) treatment. This parallels
the results of most banana field research where physiological
responses were reportedly disrupted by soil moisture levels
below 66% ASM or 33% depletion of ASM (10,40,126).
153

154
Although gas exchange processes were reduced with
decreasing irrigation levels, midday LWP in plants at W2
(20+4 ml per 150 cm3 container daily; 65 to 75% CC) were
comparable to those at the W3 level. The morphological
mechanism of lamina leaf folding, commonly seen in field
plants at midday, was evident in the present study in plants
that were water stressed to LWP of -0.51 to -0.65 MPa.
WUE has generally been quantified on a
yield/evapotranspiration basis in banana field experiments
(10,11) and though reported as leaf PS/leaf TR in this study,
there was some parallel between the two interpretations.
Plants in the greenhouse experiment grown under the W2
irrigation regime maintained PS with decreasing TR rates thus
increasing photosynthetic WUE over the W3 treated plants. At
the W1 irrigation level (10+2 ml per 150 cm3 container daily;
50 to 60% CC) plants could not effectively maintain any of
the measured physiological processes. There was chlorophyll
degradation as evidenced by leaf chlorosis and PS was
severely reduced. Although not considered drought tolerant,
the banana plant can apparently increase its photosynthetic
WUE under declining irrigation.
Under growth room conditions, effects of irrigation
volume on physiological responses were not as drastic as in
the greenhouse experiment. At midday hours, plants under the
W2 irrigation volume exhibited similar PS, CS and TR as those
at the higher irrigation level. This increase in
physiological responses at the W2 level was attributed mainly

155
to the effect of increased irradiance on stomata! opening and
PS.
Progressive reductions in physiological parameters in
ixora were also observed with decreasing irrigation volumes.
WUE however, declined in parallel with decreasing irrigation
levels and plants exhibited no apparent stress relieving
mechanisms. In the drying cycle study, leaves wilted after
irrigation was withheld for 4 days and abscised under further
stress. Under growth room conditions, decreasing irrigation
volumes did not affect physiological responses in ixora as
severely as in the greenhouse and WUE decreases were also
moderated.
Plant responses to RZTs of 28°, 33°, 38° and 43°C
imposed for 2 weeks were reported in Chapter IV. While the
33°C RZT induced maximum rates of gas exchange processes in
banana under greenhouse conditions, plants grown in the
growth room attained highest PS, CS and TR at the 38°C RZT.
All parameters were reduced by the 43°C RZT under both
environmental conditions. Most tissue-cultured banana plants
are started in black polyethylene containers and RZTs above
33°C are commonly attained in such containers (59). Results
from this study therefore indicate that nursery plants could
be grown in such containers provided that media temperatures
are limited to a maximum of 38°C.
An interesting comparison between RZT and water stress
effects was initially observed in the short-term RZT banana
experiments. Plant water status as reflected by LWP generally

156
declined with increasing RZT but plants did not exhibit leaf
folding or show any chlorosis as exhibited in the directly
water-stressed plants of the irrigation experiments. Some
mechanism of conditioning or maintaining turgor under reduced
LWP was theorized. In the subsequent long-term studies,
carbohydrate analyses suggested that there may have been
osmotic adjustment.
Measured physiological processes in ixora were not
significantly different at RZTs of 28°, 33° and 38°C but PS,
CS and TR were reduced by the 43°C RZT. PS at 38° and 43°C
was decreased compared to the lower RZTs in the growth room
experiment. This response of ixora to increasing RZT was
generally similar to that exhibited in another rubiaceaeous
genus, coffee (39), in which a RZT of 33°C was reported to be
optimum for growth and nutrient absorption.
In the final experiments, two irrigation volumes were
factorially combined with four RZTs under greenhouse and
growth room environments. Some interactions between RZT and
irrigation volume occurred that could possibly be exploited
in the container production of the two plants. Banana
exhibited tolerance to increasing RZT up to 38°C with
significantly decreased rates in measured physiological
responses induced by the 43°C RZT in the greenhouse
environment. Under growth room conditions, increased
irrigation (150+15 ml per 1200 cm3 container daily; 902 to
1002 CC) significantly increased midday PS in plants grown at
a RZT of 33°C. At 38°C RZT, overall PS declined but increased

157
irrigation application had no effect. Stress effects induced
by the 43°C RZT were moderated by increased irrigation in the
high light condition of the growth room, but increased
watering was actually detrimental to plants at the 43°C RZT
in the greenhouse experiment. Root injury at the 43°C RZT
probably caused reduced root absorptive capacity (118) and
this was aggravated by the relatively small container volume
in the greenhouse experiment.
The observed interactions could have important
implications in the nursery phase of banana production. If
growth medium temperatures are maintained at 33°C, increased
irrigation would be beneficial. At a RZT of 38°C, overall PS
would be reduced and increased irrigation would not alter
this effect. Control measures would be essential if medium
temperatures approached 43°C, since this RZT was shown to be
supraoptimal under both environmental conditions
investigated. Irrigation would need to be closely regulated
since root injury caused by the 43°C could be aggravated by
increased irrigation.
RZT stress was characterized morphologically by reduced
leaf size in banana. A distinct decrease in leaf width but
not leaf length or plant height was recorded in plants at the
38° and 43°C RZTs. A RZT-induced hormone mediated response
associated with the loss of root tips at the high RZTs was
postulated. This reducing effect on leaf area was not
overcome by increased irrigation volume, indicating another
interesting contrast between water and RZT stress effects.

158
The relatively small leaves at higher RZTs had increased
chlorophyll concentration but this was not related to
increased PS or shoot dry weight.
Carbohydrate analyses revealed significant differences
in shoot/root and sugar/starch partitioning patterns.
Comparison of the observed carbohydrate status with other
physiological parameters suggested a possible role of sugars
in allowing plants to maintain turgor under RZT-induced
decreases in LWP. More detailed analyses, however, are needed
to validate these theories and further interrelate RZT and
water stress effects. The increase in stem diameter but not
stem dry weight also supported the theory of increased plant
turgor as one response to increased RZT. Because of the
insignificant size of the corm in the plants under study,
neither dry weight nor carbohydrate determinations were made
of this tissue. This was one disadvantage of using banana
plantlets in this study, since effects on the corm can
influence final growth and yield performance of banana
plants.
In the long-term study, carbohydrate partitioning in
ixora appeared to be influenced by the RZT-induced axillary
shoot growth and environmental conditions. New shoot growth
represented an additional sink demand. However, because of
the relatively low irradiance level and thus PS in the
greenhouse, assimilates produced in the green house-grown
plants could not apparently supply both the additional shoot
growth and the increased demands of roots at increasing RZTs.

159
This led to decreased root sugar content. Supply of
assimilates in plants at the higher light level in the growth
room environment was apparently adequate for both sinks and
root sugar status was higher at the 33° and 38°C RZTs.
RZTs above 28°C caused increased axillary shoot growth
in ixora. This finding is in agreement with similar effects
of RZT in coffee (39), pepper (44) and avocado (148) but no
direct physiological explanations for the increased shoot
growth were reported. High air temperatures have also
reportedly caused non-flowering orthotropic shoots in coffee
(92) and a hormonal imbalance theory was postulated. The
induced shoot growth in ixora influenced final growth
parameters but was not accompanied by increased PS or TR.
Increased vegetative growth was apparently at the expense of
flowering and this led to the suggestion of a RZT-induced
hormonal influence, possibly cytokinin, on plant growth and
development.
Interactive effects of RZT and irrigation volumes
observed in these studies could therefore have significant
applications to the container phase of banana production
schemes that convert to the tissue-culture method of
propagation. Since ixora is used extensively as a
f1 or icultura 1 landscape plant, container medium temperatures
above 33°C in the nursery could have serious implications in
its retail value.

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BIOGRAPHICAL SKETCH
Christopher Ramcharan was born December 25, 1943, in
Port-of-Spain, Trinidad. He attended Woodbrook Presbyterian
Elementary School and St.Mary's College, Port-of-Spain,
graduating in 1961. He entered the University of the West
Indies, St.Augustine, Trinidad, in 1962 and graduated in July
1965 with the BSc in Tropical Agriculture. From 1965 to 1972
he was employed with the Department of Agriculture and served
as Agricultural Officer at the Central Experiment Research
Station, Centeno, and Superintendent of the Royal Botanic
Gardens in Port-of-Spain, Trinidad.
In 1972, he was awarded a Trinidad and Tobago
Government Development Scholarship which enabled him to enter
the University of Florida and earn a MS degree in ornamental
horticulture in December 1975. From 1978 he has been on the
staff of the Virgin Islands Agriculture Experiment Station as
research horticulturist. In August 1984 he re-entered the the
University of Florida as a joint investigator of heat stress
of tropical fruits and ornamental plants on a USDA Tropical
Agriculture 406 Grant. During this time he also pursued
graduate work towards the PhD degree in the area of root-zone
temperature stress physiology.
172

I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality, as
a dissertation for the degree of Doctor of Philosophy.
Mftr
hairman
Dewayne ¿ingram,
Associate Professor of
Horticultural Science
I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality, as
a dissertation for the degree of Doctor of Philosophy.
James Barrett
Associate Professor of
Horticultural Science
I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality, as
a dissertation for the degree of Doctor of Philosophy.
Associate Professor of
Agronomy
I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality, as
a dissertation for the degree of Doctor of Philosophy.
AÍ
•ril Nell
Associate Professor of
Horticultural Science
V

I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality, as
a dissertation for the degree of Doctor of Philosophy.
Will t/am Wi I tbank
Professor of
Horticultural Science
This dissertation was submitted to the Graduate Faculty of
the College of Agriculture and to the Graduate School and was
accepted as partial fulfillment of the requirements for the
degree of Doctor of Philosophy.
May 1987
Agriculture
Dean, Graduate School

UNIVERSITY OF FLORIDA
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