Front Cover
 Title Page

Group Title: Vanguard (East Lansing, Mich.)
Title: Temperature X photoperiod, adaptation and yield in phaseolus vulgaris
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00086755/00001
 Material Information
Title: Temperature X photoperiod, adaptation and yield in phaseolus vulgaris
Series Title: Temperature X photoperiod, adaptation and yield in phaseolus vulgaris
Uniform Title: Vanguard (East Lansing, Mich.)
Physical Description: 11 p. : col. ill. ; 23 cm.
Language: English
Creator: Wallace, D. H ( Don H )
Masaya, Porfirio N., 1935-
Gniffke, Paul A ( Paul Arthur ), 1947-
Bean/Cowpea Collaborative Research Support Program
Publisher: Bean/Cowpea CRSP, Michigan State University
Place of Publication: East Lansing
Publication Date: 1984
Subject: Common bean -- Climatic factors -- Guatemala   ( lcsh )
Common bean -- Climatic factors -- Ecuador   ( lcsh )
Common bean -- Growth -- Tropical conditions   ( lcsh )
Genre: non-fiction   ( marcgt )
Spatial Coverage: Guatemala
Funding: Funded through USAID/BIFAD Grant No.
Statement of Responsibility: Donald H. Wallace, Porfirio N. Masaya, Paul A. Gniffke.
General Note: "Vanguard (East Lansing, Mich.), Volume 1, no. 1, March 1984."
 Record Information
Bibliographic ID: UF00086755
Volume ID: VID00001
Source Institution: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: oclc - 234307156

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Full Text

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Michigan State University

The Bean/Cowpea CRSP is a program of coordi-
nated projects in Africa and Latin America address-
ing hunger and malnutrition through research on the
production and utilization of beans (Phaseolus vul-
garis) and cowpeas (Vigna unguiculata). The goal of
the Bean/Cowpea CRSP reflects the Title XII "Famine
Prevention and Freedom from Hunger" mission of
the US Foreign Assistance Act under which the pro-
gram is funded. The goal is to establish active and
vigorous collaborative research efforts that will con-
tribute to the alleviation of hunger and malnutrition
in developing countries by improving the availability
and utilization of these foods. In the true spirit of
collaboration, the CRSP also makes a significant con-
tribution to agriculture in the US through the in-
creased knowledge and materials generated by the
research partnerships with Host Countries (HCs).
The research findings and identified biological re-
sources hold potential for solving or reducing impor-
tant agricultural constraints to bean and cowpea
production and consumption in legume producing

Beans and cowpeas are dietary staples in the HCs
associated with this CRSP. Among many families,
these legumes provide the major source of high
quality, affordable protein as well as an important
source of B vitamins. Beans and cowpeas generally
are grown as food for household consumption,
rather than as export crops. They are typically grown
on subsistence farms and, in some countries, are
grown solely by women, on whose shoulders rests
the major responsibility for providing the food for
family consumption. CRSP research seeks to
strengthen the resources available to these produc-

To support its goal, the CRSP directs its attention

1. Building strong and collegial professional rela-
tionships among the HC and US researchers in
each project;

2. Making financial resources available for both
HC and US research activity;

3. Emphasizing multidisciplinary research inte-
grating production and non-production issues;

4. Focusing on research in traditional settings;

5. Paying specific attention to the roles and par-
ticipation of women;

6. Being alert to mechanisms for information dis-
semination; and

7. Supporting HC educational opportunities to
strengthen long-term legume research capabil-
ity within the country.

CRSP research is concerned with genetics and
plant breeding, agronomics, economics, nutrition
and socio-cultural factors. A global research plan,
developed jointly by HC and US colleagues, formed
the basis for the design of the collaborative projects.
These vigorous international research partnerships
directly involve research institutions in thirteen HCs,
two international centers and fourteen US agricul-
tural research institutions, which include the nine
having lead roles in the CRSP.

In making available to the international agricultural
research and development system an avenue to the
US agricultural research network, the Bean/Cowpea
CRSP is making important contributions to the re-
solution of difficult and persistent problems as-
sociated with bean and cowpea production and utili-
zation. Across the many participating disciplines and
nations, there is emerging from this CRSP a
heterogenous cadre of professional research col-
leagues prepared to share the unique resources of
one another in addressing long-term troublesome
constraints to adequate food and nutrition in poverty
regions of the world. The CRSP Vanguard Series is a
publication of the important research findings which
are evolving from these collaborative professional re-

Temperature X Photoperiod,

Adaptation and Yield in

Phaseolus vulgaris

Dr. Donald H. Wallace
Department of Plant Breeding and Biometry
Cornell University

Dr. Porfirio N. Masaya
Bean Program Leader

Paul A. Gniffke, PhD Candidate
Department of Plant Breeding and Biometry
Cornell University

VANGUARD, Vol. 1, No. 1
Ardeshir Ghaderi, Editor
Bean/Cowpea CRSP
Michigan State University
March 1984

Funded through USAID/BIPAD Grant No. AID/DSAN-XII-G-0261

Temperature x Photoperiod,

Adaptation and Yield in

Phaseolus vulgaris

Extensive variations exist from one location to another in the amount
of rainfall and the duration of the rainy seasons) within almost all
tropical countries. High yield of quality beans requires sufficient rainfall,
with maturity occurring during a rainfree period. Beans cannot be
grown in dry seasons without irrigation or residual moisture, and most
plantings depend on natural rainfall. For these reasons, the duration of
growth, i.e., days from planting to seed maturity, is an important
feature that adapts a cultivar to a geographical locality. This includes
temperate locations where temperatures determine the growing season
duration. Every location tends to have a specific duration of active
growth that makes for maximum yield. Besides the above mentioned
water and temperature relationships, the choice of crops to be grown in
multiple cropping with beans can alter the required days to maturity.
Also, mean and absolute day and night temperature can greatly modify
the duration of growth required for a given bean cultivar. Under each
temperature regime, yields are generally increased per unit land area as
the duration of growth is extended. The duration may be 60 days in
warm localities and up to about six months in high elevation tropical
locations. Simultaneously with increased growth duration, yield per
land area per day may be drastically reduced. Therefore, total yield
may be maximized with tandem plantings of early maturing cultivars.
For all these reasons, breeding for maturity, adaptation and yield are
critical objectives of bean improvement programs in Guatemala and
Ecuador, the project sites of the research reported here, but also in all
tropical and temperate countries.
Beans express a response to long photoperiods ranging from
complete insensitivity in which photoperiod has no effect on timing
or intensity of flowering, to different degrees of quantitative sensitivity -
in which flowering is delayed to some extent by long photoperiods,
to nearly qualitative sensitivity in which flowering seems to be
indefinitely postponed by long days. A variety's sensitivity to
photoperiod may have a critical impact on its usefulness in many
growing locations particularly in mid to high latitudes. A variety's
relative temperature adaptation vis-a-vis a location's growing
temperatures also has great influence on flowering time and time to
maturity and may interact strongly with photoperiod response.

Three flowering responses to temperature occur in the field and in
controlled environments:
1. The below-optimum temperature for flowering response of the
given cultivar (genotype). This response is a change toward earlier
flowering as the temperature is raised.
2. The optimum temperature for flowering as indicated by the
minimum expressed days to flowering of each genotype. This
minimum is called the flowering tendency.
3. The above-optimum temperature for flowering response of each
genotype. This response is a change toward later flowering as the
temperature is raised. These three responses to changes of
temperature, with daylength remaining constant, plus simultaneous
direct and interaction responses of flowering to changes of
photoperiod, collectively make up three major physiological-genetic
components of days to flower and consequent maturity, adaptation
and yield:

a. The photoperiod x below-optimum temperature response.
b. The flowering tendency as expressed at the optimum
c. The photoperiod x above-optimum temperature response.

The interaction of the varied daylength-temperatures of different
geographical locations divides bean production of the world into areas
where above-optimum temperature, optimum temperature, and
below-optimum temperature for flowering, in combination with short
vs. moderate vs. long daylengths, contribute differentially to
quantitative delays in days of flowering. The delay is beyond that
minimal days of the flowering tendency of each genotype which is
expressed only at the optimal photoperiod-temperature for the
genotype, i.e., usually at 800-1000 meters in the tropics.
There is evidence for genetic differences among bean cultivars with
regard to response to daylength and temperature. CRSP-sponsored
research at CIAT has shown that electric-light extended, long days delay
flowering of bean genotypes most strongly at high temperatures (low
elevations), intermediately at intermediate temperatures (moderate
elevations) and little or not at all at low mean temperatures (high
elevations). This agrees with our observation in Guatemala that,
contrary to possible expectation, the delay of time of flowering as
temperature is lowered is attended by a lowering of node number to
flower, which would tend to give earlier flowering; the change of node
number is more closely correlated with temperature than with days to

Other research at CIAT has shown that, at continuous moderate
(near optimal) temperature, an extension of daylength to eighteen
hours delayed flowering of a sensitive cultivar by seven days and
increased yield by 50 and 72% in two plantings. The same
photoperiod-temperature-caused delay of flowering in temperate zones
is likely to decrease yield because pod development in this case takes
place during the lower fall temperatures.
International yield and adaptation nurseries of CIAT show flowering
of photoperiod-temperature sensitive bean cultivars to be increasingly
delayed at locations with longer and longer daylengths. Our research
shows this photoperiod-modulated delay to be further increased by
high temperature. This occurs because the delay is always
simultaneously influenced by both daylength and temperature.
Differential cultivar adaptation to different world areas arises in part
because genotype-directed potential for delay in node to flower (the
second process) is always modulated simultaneously and
synergistically by temperature and daylength. Number of nodes
initiated per day (the first process) is only modulated by temperature.
An optimum temperature for flowering exists for each daylength and
genotype. There is also an optimum photoperiod-temperature where
days to flower is fewer than under any other daylength-temperature.
From this optimum, all rising or lowering of temperature and
extensions of daylength delay flowering.
The above described biology of maturity, components of time of
flowering and responses of flowering to daylength and temperature
divides bean production into zones where cultivar adaptation is
achieved differently.

Regulation of Bean Maturity in Different Areas
of the World

1. Hot Lowland Tropics. It is predominantly the above-optimum
temperature for flowering component of the above-optimum
temperature x photoperiod response that limits bean adaptation and
yield in short-daylength (presumed non delaying), high temperature
tropical locations below 800 to 1000 meters. At such
above-optimum temperature locations, the change in days to flower
is a delay when there is a rise of temperature, and earlier flowering
when temperature is lowered.

2. Temperate Locations. As long recognized, daylength plus a cultivar's
genotypic sensitivity to it predominantly control the time of
flowering and maturity in long-day temperate locations. This control

Dr. Donald H. Wallace, US Principal
Investigator, Cornell University.

ICTA technician evaluating and recording
project intercropping trials, Guatemala.

Dr. Porfirio N. Masaya, Host Country
Principal Investigator, ICTA, Guatemala.

Project field trials, ICTA, Guatemala.

ICTA field hands weeding bean plots, Guatemala.

Dr. Masaya at crossing block, ICTA, Guatemala.

is by precisely the same physiological-genetic process that controls
time of flowering and maturity in lowland tropical locations. The
only difference is that in a temperate area the days to flowering are
delayed largely by the long daylength (or at least can be) rather than
by high temperature. The "can be" is because, depending on
genotypic sensitivity to photoperiod and consequently to
above-optimum temperature, a moderate above-optimum
temperature may result in a larger delay in flowering in conjunction
with a long (temperate) daylength than will a very high temperature
under a short (tropical) day.

3. Highland Tropics. Relatively few beans are grown at tropical
elevations much lower (warmer) than the main bean research
stations of ICTA in Guatemala (900 meters, Jutiapa) and of CIAT in
Colombia (1000 meters, Palmira). Both stations are located at near
optimal temperatures, i.e., at that mean temperature which gives
the minimal days to flower for most cultivars. Seventy-five per cent
of beans are grown at elevations higher than those with the
23-25C that constitute the optimal temperature for flowering.
Besides genotype, therefore, the main environmental factor
controlling maturity and consequent adaptation and yield of 75% of
tropical bean production is the below-optimum temperature for
flowering. The short daylength and low temperature both minimize
any photoperiod effect.

4. Optimal Photoperiod-Temperature in the Tropics. Such an optimum
or near-optimum photoperiod-temperature is a necessary
environment for expression of the flowering tendency of a cultivar.
The genetic variation of the flowering tendency is the major
physiological-genetic component of variable maturity at near
optimal photoperiod-temperature, i.e., at tropical elevations of 800
to 1000 meters.

Influence of Diurnal Temperature Difference

Growth chamber studies show that a small difference between day
and night temperatures minimizes the delay in flowering caused by
long daylength plus above-optimum temperature for flowering, while a
large difference amplifies the delay. Under short daylength, plus near
optimal temperature, the large diurnal difference delays flowering by a
mechanism other than abortion of flower buds that are only
microscopically visible. The flower buds develop to a visible size and
then abort a week or a day or two before anthesis, or a day or two

after anthesis at the pod setting stage. Thus, the diurnal difference
modulates the stage of flower development at which abortion occurs.

Genetics of Maturity and Adaptation

Existing knowledge about inheritance of bean sensitivity to
photoperiod indicates regulation by only a few genes. Studies have not
adequately accounted for the simultaneous above-optimum
temperature modulations through the delay in node to flowering and
through the nodes developed per day, nor as through the modulation
by photoperiod.
CRSP-sponsored research at CIAT has demonstrated that
genotype-directed, very strong, strong, intermediate, weak, and very
weak sensitivities to photoperiod-temperature can be genetically
associated with each of short, intermediate and long durations of the
flowering tendency. No inheritance studies have clearly differentiated
photoperiod sensitivity from the other physiological genetic
components of maturity.
Optimal temperatures near 18 C for some highland cultivars and
near 23-25 C for most cultivars have been shown by CRSP and CIAT
research. Thus, genetic variability in optimal temperature for flowering
exists and can be useful in breeding for cultivar adaptation. Knowledge
is needed about genetics and inheritance of the flowering tendency
which is expressed only at an optimal temperature. Besides being the
only component of maturity expressed at optimal temperatures
(moderate elevations), the minimum number of days during which a
plant has a tendency to flower is the basic number of days to flowering
to which all delays caused by daylength and/or temperature are added.

Applications to Breeding

Because below-optimum temperature for flowering is the major
controlling environmental factor for bean production in the tropics and
because every environmental factor functions through one or more of
the four identified genetic systems, research on these
physiological-genetic factors is essential to improving efficiency of
breeding for maturity, for consequent adaptation and for yield of
tropical bean cultivars. Such knowledge is most deficient for breeding
cultivars adapted to the highlands.
It is necessary to identify existing genetic variation in optimal

temperature for flowering and learn about its inheritance. How does
the effect of this genetically directed optimal temperature for flowering
of a cultivar differ for low, moderate and high elevation locations? How
does it differ for short, moderate and long day locations? What
flowering tendency and what optimal temperature will give maximum
adaptation and yield for the different elevations? Will a long or short
duration flowering tendency combined with maximal insensitivity to
photoperiod-temperature maximize yield in the lowland tropics? in
temperate zones?
Use of genetic variation in flowering tendency should be tested for
breeding of improved cultivars for both the lowland tropics and
temperate zones, as well as for moderate elevation tropics where it is
the only genetic variability that will be expressed. Depending on rainy
season duration in the tropics and on fall temperatures in temperate
areas, a long duration flowering potential with minimal possible
sensitivity to photoperiod and above-optimum temperature would
seem to be preferable over a cultivar that has its moderate or late
maturity established through its gene-directed sensitivity to
photoperiod-temperature. Proper flowering potential and minimal
photoperiod-temperature sensitivity could facilitate bean production at
low elevation, hot tropical locations. It could also minimize the large
fluctuations in bean yields that occur because of year to year
temperature variations in temperate locations.

Agronomic, Sociological and Genetic Aspects
of Bean Yield and Adaptation

Dr. Donald H. Wallace US Principal
Cornell University
Dr. Porfirio N. Masaya HC Principal
Institute de Ciencias y Tecnologia Agricola

Project Objectives

1. To determine the biological, environmen-
tal, economic and social roles of bean
production in the farming systems of
small farms.

2. To develop methods of determining the
merit of potentially useful bean produc-
tion practices using research conducted
in the environment and under the man-
agement system of small farms.

3. To determine the competitive advantages
of specific crops in bean intercropping
systems and determine the pattern pro-
viding the highest return from the appli-
cation of limited resources such as fer-

Collaborating researchers
Dr. Patricia Garrett, Cornell University
Dr. Roger F. Sansted, Cornell University
Ing. Selvin Arriaga, ICTA

4. To develop credible procedures for
measuring (or estimating) the degree of
acceptance of new practices by small
farmers and determine the merit of
feedback from such procedures for es-
tablishing research objectives.

In this project the socio-agronomic bases
are sought for bean farming systems of high-
land Indians. Change from monocropped
bush, to bush with corn, to climbing bean with
corn occurs as elevation increases. The human
and ecological bases for the maintenance of
one system over the others are investigated.
Two years of field research in bean physiology
has been conducted at different elevations
under natural 12 hour and 18 hour electric
light extended daylengths. This research, the
focus of the present report, was based in part
on the previous years' growth chamber re-
search findings. Cultivator adaptation compari-
sons between temperate and tropical zones
have also been studied.

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