Ann.Bot.

51, 531-543, 1983

A Quantitative Model of Reproductive

Development in Cowpea [Vigna unguiculata (L)
Walp.] in relation to Photoperiod and Temperature,
and Implications for Screening Germplasm
P. HADLEY, E. H. ROBERTS, R. J. SUMMERFIELD
University of Reading, Plant Environment Laboratory, Shinfield Grange, Cutbush Lane, Shinfield,

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Reading, Berks RG2 9AD

and F. R. MINCHIN
Plant and Crop Physiology Department, Grassland Research Institute, Hurley, Maidenhead, Berks SL6 5LR

Accepted: 26 September 1982

ABSTRACT
Factorial combinations of four photoperiods (10 h, 11 h 40min, 13 h 20min and 15 h) and three night
temperatures (14, 19 and 24 C) combined with a single day temperature (30 C) were imposed on nodulated
plants of 11 cowpea accessions [Vigna unguiculata (L) Walp.] grown in pots in growth cabinets. The times to
first appearance of flower buds, open flowers and mature pods were recorded. Linear relationships were
established between the reciprocal of the times taken to flower and both mean diurnal temperature and
photoperiod. When the equations describing these two responses are solved, the time to flower in any given
photothermal regime is predicted by whichever solution calls for the greater delay inflowering.Thus in different
circumstances flowering is controlled exclusively by either mean temperature or photoperiod. The value of the
critical photoperiod is temperature-dependent and a further equation, derived from the first two, predicts this
relationship. Considered together as a quantitative model these relationships suggest simple field methods for
screening genotypes to determine photo-thermal response surfaces.

Key words: Vigna unguiculata (L) Walp., cowpea, reproductive development, photoperiod, temperature,
germplasm.

INTRODUCTION

There is no doubt that the rate of reproductive development, yield potential and actual
seed yield in cowpea [Vigna unguiculata (L) Walp.] are extremely sensitive to weather
and climate (e.g. Summerfield et al, 1982). Since it is impossible to modify seasonal
changes in the photo-thermal environment in the field, it is necessary to fit the
developmental responses of cultivars to the environments which prevail in different
ecological zones. A greater understanding is needed not only of the genetic factors which
control crop duration but also of the environmental factors that react with them (Aitken,
1974). From previous research on cowpea in controlled environments and in the field
(e.g. Huxley and Summerfield, 1974), almost 90 genotypes have been classified according
to their sensitivity to photoperiod and night temperature. However, the original emphasis
focused intentionally on the relatively narrow range of conditions which prevail in the
lowland humid tropics (i.e. within 7 N or S of the equator) and did not extend to the
widest range of photo-thermal conditions experienced by cowpea crops in West Africa.
In the work reported in this paper we selected 11 accessions known to respond
differently to the small ranges of photoperiod and night temperature tested hitherto, and
we now describe their responses to a much wider range of each of these factors.
0305-7364/83/040531 + 13 $03.00/0 1983 Annals of Botany Company
532 Hadley et al.Reproductive Development in Cowpea

MATERIALS AND METHODS

Environmental simulation
Factorial combinations of four photoperiods (10 h, 11 h 40 min, 13 h 20 min and 15 h)
and three night temperatures (14, 19 and 24 C, all 0-5 C) were imposed on nodulated
plants of 11 cowpea accessions [V. unguiculata (L) Walp.] grown in pots in controlled
environment growth cabinets. A mixture of 95 per cent 'Daylight' fluorescent tubes and
5 per cent tungsten lighting (according to the total nominal wattage per cabinet) provided
a radiation flux density of 164Jm" ! s~ 1 (400-700 nm) at pot level. This would be
approximately equivalent in terms of photosynthetically-active radiation in daylight
(300-3000 nm) to 11-8, 13-8,15-8 and 17-8 MJm~a in the four photoperiods investigated.

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Photoperiods and night temperatures were selected to cover the ranges likely to be
experienced by cowpea crops grown at various latitudes and altitudes in the humid and
the semi-humid tropics. Night temperature rather than day temperature was varied as
our previous work had suggested that this factor was more important in controlling
flowering. The scale of each factor was chosen to form an equally spaced series:
successively longer photoperiods differed by 100 min and successively warmer nights by
5C. Day temperature (300-5C), relative humidity (702-5 per cent) and CO 8
concentration (340 15 ppm) were maintained at a constant level in each cabinet.
When investigating factorial combinations of photoperiod with either day or night
temperature, confounding of some sort is inevitable, and any experimental design
involves some sort of compromise. In these experiments we chose to use artificial
environments which attempt to approach natural conditions, and so photoperiods were
not differentiated using low intensity lighting but were confounded with total radiation.
Thus, plants grown in the 11 h 40 min, 13 h 20 min and 15 h photoperiods received 16-9,
33-9 and 50-8 per cent more photosynthetically-active radiation, respectively, than those
grown in the 10 h regime. Furthermore, in natural conditions the diurnal change in air
temperature lags asymmetrically behind that of radiation, so day temperatures in the
cabinets were set to start 2 h after lights came on and to terminate at 'lights off'.
Consequently, as photoperiod increased from 10 h to 11 h 40 min, 13 h 20 min or 15 h,
daytime temperature sums (integral of time and temperature) were increased, as were
mean diurnal temperatures although to a lesser extent since night temperature sums
became progressively smaller. A useful feature of this design is that mean diurnal
temperatures were different in all 12 environments used (Table 1).

Plant husbandry and culture

Techniques were based on those developed for tropical grain legumes and described in
detail elsewhere (Huxley and Summerfield, 1976; Summerfield, Huxley and Minchin,
1977). Seeds of all accessions were weighed individually and only those from the median
range of each stock were used. Batches of seeds were surface sterilized for 2 min in 0-2
per cent mercuric chloride in 001 M HC1, washed three times in sterile distilled water,
blotted dry, reinoculated with strain of Rhizobium CB 756 (see details in Summerfield
et al., 1977) and sown three to each 18 cm diameter plastic pot containing a mixture of
vermiculite, sand, gravel and loamless peat compost without fertilizer (4:2:4:1 by
volume) and soaked with deionized water. After sowing, the seeds were covered with a
2 cm layer of washed, dried granite grit.
Seedlings were given only deionized water until their first pair of true leaves was fully
expanded. Thereafter, sufficient nutrient solution, complete except for containing only
20 mg I"1 inorganic nitrogen (see Summerfield et al., 1977), was delivered automatically
two to four times each day so that drainage through the pots occurred at every
 Hadley et al.Reproductive Development in Cowpea 533

TABLE 1. Day, night and mean diurnal temperature (C) and total temperature sums
(centigrade hours above 0 C) in each photo-thermal regime

Day-night Temperature sum (Cxh) Mean diurnal

Daylength temperature temperature
(h) (C) Day* Nightf Total (Qt
10 30-14 268 1% 464 19-3
10 30-19 278 266 544 22-7
10 30-24 288 336 624 260
11 h 40min 30-14 318 173 491 20-5
11 h 40 min 30-19 . 328 234 562 23-4

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11 h 40 min 30-24 338 296 634 26-4
13 h 20 min 30-14 368 149 517 21-5
13 h 20 min 3O-19 378 203 581 24-2
13 h 20 min 20-24 388 256 644 26-8
15 30-14 418 126 544 22-7
15 30-19 428 171 599 250
15 30-24 438 216 654 27-2

* Since the switch to the nominal day temperature was delayed for 2 h after the start of the photoperiod,
these values are calculated as: [(Duration of photopenod 2) x nominal day temperature+ (2 x nominal night
temperature)].
t Calculated as: (Duration of dark period x nominal night temperature).
X Total temperature sum divided by 24.
Environments used in previous investigations (e.g. Huxley and Summerfield, 1974).

application. At this concentration of applied nitrogen cowpea plants make the most
efficient use of dinitrogen fixation (Summerfield et al., 1982).
Six replicate pots of each accession were sown in each environment. Seedlings were
thinned after 14 days to leave single uniform plants in each pot. Ten days later the plants
were regraded and three replicates of each accession were discarded in order to avoid
overcrowding in the cabinets.
Two experiments were carried out, the first starting on 3 August 1978 and the second
on 9 August 1979. In both cases eight accessions were investigated, five of which were
common to both experiments. Accession numbers are those of the International Institute
of Tropical Agriculture, Ibadan, Nigeria. Those common to both experiments, with their
origin, where known, in parentheses, were: TVu 3 (Nigeria), TVu 317 (USA), TVu 465-1-2
(Nigeria), TVu 1009 (USA) and TVu 1503 (USA); those investigated in the first
experiment only were: TVu 167, TVu 946-1-1 (Nigeria) and TVu 1188 (Uganda); and
those included in the second experiment only were: TVu 136 (USA), TVu 1047 (USA)
and TVu 3699 (Nigeria). Records were made on individual plants of the respective times
(days) from sowing to (a) emergence, (b) appearance of first flower bud (distinguishable
from vegetative buds by the naked eye), (c) appearance of first open flower (corolla colour
visible) and (d) maturity of first pod (judged by changes in colour, texture and
succulence).
Since those five accessions which were grown in both experiments did not differ in any
phenological expression by more than 2 days, results from both experiments have been
combined.
RESULTS
Irrespective of genotype, 90 per cent emergence occurred between 4 and 6 days after sowing
in all environments. Thus, the temporal differences in subsequent observations were
entirely due to differences in rates of reproductive development between genotypes or
534 Hadley et al.Reproductive Development in Cowpea
environments. Almost all replicates of a given genotypefloweredon the same day in any
one photo-thermal regime; the maximum variation was typically 2 days.
Certain accessions (e.g. TVu 3 and TVu 3699) were acutely sensitive to photopenod
and showed no sign of initiatingflowerbuds in some treatment combinations - even after
3 months from sowing when the experiments were terminated (Fig. 1).
(a) (b)
100

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100

80 V 1 A; Al
J
A
ni
a
Al A A A ?4 At Ai

60
\ \
Rp Rp Rp
40
19 21 23 25 27 19 21 23 25 27 19 21 23 25 27
Mean diurnal temperature (C)

FIG. 1. Effects of photo-thermal regime on successive stages of reproductive development of each

of three cowpea genotypes acutely sensitive to photoperiod: (a) TVu 167, (b) TVu 3, (c) TVu 3699.
Photoperiods are denoted by O, 10 h; Q, 11 h40min; A, 13 h 20min; , 15 h. The symbols with
attached arrows pointing upwards at 95 days ( ) denote that plants had not reached the relevant
stage of development when the experiment was terminated at this time. Solid regression lines were
calculated according to Eqn (1) and independently for each cultivar. Broken regression lines were
calculated after constraining each regression to a common base temperature (r0) of 8 C as in Fig.
2. Vb,f and Rp denote first visible buds,firstflowersand first ripe pods, respectively.

Photoperiod-insensitive cultivars
Six genotypes responded similarly and produced buds andfloweredsoonest (or almost
so) in the longest day-warmest night (15 h-24 C) treatment combination and, conversely,
remained vegetative for the longest time in the shortest day-coolest night (10 h14 C)
regime (Fig. 2). It is clear from an example (TVu 1009) chosen from this group [Fig. 3(a)]
that flowering occurred earlier the longer the photoperiod and the warmer the night
temperature. However, accessions of this type are not quantitative long day plants
sensitive to night temperature as Fig. 3(a) suggests, since variations in flowering time
 Hadley et al.Reproductive Development in Cowpea 535
100

80 -

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20

19 20 21 22 23 24 25 26 27 19 20 21 22 23 2425 26 27
FIG. 2. Effects of mean diurnal temperature on successive stages ofreproductivedevelopment of each
of six photoperiod-insensitive cowpea genotypes: (a) TVu 176, (b) TVu 317, (c) TVu 465, (d) TVu
946, (e) TVu 1009, (0 TVu 1503. Days from sowing to particular stages of development are denoted
by O,firstvisible buds; Q.firstopenflowers;A,firstripepods. Solidregressionlines were calculated
according to Eqn (1) and independently for each cultivar. Broken regression lines were calculated
after constraining eachregressionto a common base temperature (t0) of 8 C (e.g. forfirstflowering,
1//"= 0 at 8 Q.

can be explained with great precision and simplicity as a function of mean diurnal
temperature in all environments [Fig. 3(b)]. The apparent photoperiodic and night
temperature responses are due to confounding between daylength and mean temperature
in the treatments imposed (Table 1). The genotypes shown in Fig. 2 are completely
insensitive to photoperiod; their time to flowering is simply related to mean temperature
in a manner described by Eqn (1), in which /denotes the number of days from sowing
to the onset of flowering, / i s mean temperature and a and b are constants:
\lf=a+bi. (1)
Thus, the rate of progress towards flowering (the reciprocal of the time taken) is a
linear function of mean temperature. If Eqn (1) applies, a base temperature, t0, can be
defined as the temperature at which the rate of progress towards flowering is zero (i.e.
536 Hadley et al.Reproductive Development in Cowpea

100
? 90
) 80
70

\ 60

A > 55
50 /
\
45
40

35

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-^ >^ii 20
140 ^^

FIG. 3. Effects of photo-thermal regime on flowering in cowpea accession TVu 1009. Response surfaces
show (a) effects of photoperiod and night temperature on days from sowing to flowering (f) and (b)
effects of photoperiod and mean diurnal temperature on days from sowing to flowering (/"and I If
on right and left hand ordinates, respectively). Solid circles denote the experimental mean values in
each of 12 environments (Table 1).

1 // = 0), and its value can be calculated from the constants of the regression Eqn (1) using
the expression: ^ = __ajb (2)

Furthermore, the thermal sum, 7} (in day degrees above the base temperature), required
for flowering is given by: -, _ , ,, ,,.
if - i/b. (3)
As for the appearance of first open flowers, the times to appearance of buds visible
with the naked eye and, subsequently, of the first pods to ripen on these photoperiod-
insensitive genotypes were also inversely related to mean diurnal temperature (Fig. 2).
The values of the constants derived from the regressions involving all three successive
stages of reproductive development are given in Table 2.

Photoperiod-sensitive cultivars
Five accessions (Figs 1 and 4) responded to photoperiod in a manner typical of
quantitative short-day plants (Vince-Prue, 1975). Three of them were so acutely sensitive
to longer photoperiods that they did not come into flower within 95 days from sowing
in photoperiods of 13 h 20 min (TVu 3 and TVu 3699) or 15 h (TVu 167), even though
they had flowered as early as 40 to 41 days from sowing in warm, photo-inductive
environments. Similarly, a combination of very short days (10 h) and the warmest nights
(24 C) was most inductive for the two remaining genotypes (TVu 1047 and TVu 1188)
in this group, but these were less sensitive to photoperiod and they flowered in all
less-inductive regimes within 95 days. A typical example of the responses of genotypes
in this group is presented for TVu 1188 in Fig. 5.
Again, since night temperature per se was thought originally to be an important factor
in the environmental regulation of reproductive development in cowpeas (e.g. Huxley
and Summerfield, 1974) the data for TVu 1188 were plotted against both this factor and
photoperiod [Fig. 5(a)]. When presented in this way the data suggest a complex
interaction between these two factors but, as Fig. 5(b) shows, when the results are plotted
 Hadley et al.Reproductive Development in Cowpea 537
TABLE 2. Values of constants in Eqn (1) derived from regressions of respective inverse
durations (days from sowing) for first visible buds, first open flowers and first ripe pods
against mean diurnal temperature for each of six photoperiod-insensitive genotypes of
cowpea

Standard Base
error temperature
Genotype Intercept (a) Slope (b) of slope (Q (~alb) r1

First visible buds

TVu 176 -00390 000338 000012 11-5 0-962
TVu 317 -00045 000150 000009 30 0-729
TVu 465 -00107 000180 000009 5-9 0-850

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TVu 946 -00101 000194 000016 12 2 0-804
TVu 1009 -00238 000262 000010 91 0-905
TVu 1503 -00129 0-00240 000014 5-4 0-812
First open flowers
TVu 176 -00200 0-00191 000005 10-5 0-979
TVu 317 -00108 000134 000004 81 0-937
TVu 465 -00156 000161 000005 9-7 0-940
TVu 946 -00132 000160 000007 8-3 0-940
TVu 1009 -00193 000179 000004 10-8 0-963
TVu 1503 -00153 000176 000004 8-7 0-961
First ripe pods
TVu 176 -00110 000112 000009 9-8 0-802
TVu 317 -00053 000081 000003 6-5 0-919
TVu 465 -00104 0-00107 000007 9-7 0-788
TVu 946 -00037 0-00086 000007 4-3 0-801
TVu 1009 -00064 000089 000006 7-2 0-774
TVu 1503 -00096 000110 000003 8-7 0-935

as the reciprocal of time to flowering against mean diurnal temperature and photoperiod,
a relatively simple picture emerges.
In short (inductive) photoperiods this accession shows the temperature response
typical of photoperiod-insensitive genotypes [Fig. 3(b)], as described by Eqn (1).
However, a second response, superimposed on the temperature response surface, shows
a progressive delay in the onset of flowering as photoperiod increases [Fig. 5(b)]. The
response to photoperiod can be described by a similar Eqn (4) to that used earlier for
temperature, and in which p denotes photoperiod and a' and b' are constants (and b'
is always negative for short day plants):
l/f=a' + b'p. (4)
The effects of photoperiod and temperature on successive stages of reproductive
development in these photoperiod-sensitive accessions are shown in Figs 3 and 4.
Clearly, each of these photoperiod-sensitive accessions has a similar response to mean
temperature when grown in short (inductive) photoperiods (Eqn 1) and their response
to longer days is equally well explained by Eqn (4). Thus, flowering in these genotypes
is determined exclusively by either photoperiod or mean temperature, whichever causes
the greater delay. Therefore, in any photo-thermal environment,/can be predicted by
solving Eqns (1) and (4) and the correct solution is that which indicates the longer period
to flowering.
The critical photoperiod, pc, defined here as that photoperiod which, if exceeded, causes
a delay in flowering, is represented by the line of intersection of the response surfaces
for photoperiod and mean temperature [Fig. 5(b)]. Thus, algebraically, pc can be
538 Hadley et al.Reproductive Development in Cowpea

(a) (b)
100

80

60

40

20

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100

80

60

40

; ; i i
80
A
60 -^00
Rp
19 20 2122 23 24 25 2627 28 192021 22 23 24 25 26 27 28
Mean diurnal temperature (*C)
Fio. 4. Effects of photo-thermal regime on successive stages of reproductive development of each of
two cowpea genotypes moderately sensitive to photoperiod: (a) TVu 1188, (b) TVu 1047. Explanation
of symbols as in Fig. 1. Solid horizontal lines, which indicate delays from the temperature curves
attributable to photoperiod, were fitted by eye.

determined when the values for/, given by solving Eqns (1) and (4), are identical. In other
words:
P=PC when a + bi= a' + b'p
and so, = (a-a' + bt)/b'. (5)
It is clear from Fig. 5(b) that critical photoperiod varies with mean temperature and,
for TVu 1188 over the range of temperatures investigated here (Table 1), it changed by
3 h 35 min, i.e. a decrease of about 27 min per C increase in mean temperature. For
genotypes acutely sensitive to photoperiod (e.g. TVu 3 or TVu 3699; Fig. 1) there was
little or no detectable effect of temperature on critical photoperiod. In both these
genotypes, as in most others, the onset of flowering in photoperiods shorter than 11 h
40 min was determined solely by mean temperature, but in photoperiods of 13 h 20 min
or longer, flowering was delayed so markedly that it.did not occur within the duration
of the experiment. For these two genotypes the slope of the photoperiod response surface
must be very steep (i.e. b' is very large) and so, as Eqn (5) predicts, pc will vary only
slightly with temperature.
Photo-thermal effects on the durations tofirstvisible buds and then ripe pods followed
a substantially similar pattern to those for first open flowers (Fig. 4). However, because
 Hadley et al.Reproductive Development in Cowpea 539

75

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''Cj 2 8 10,

Fio. 5. Effects of photo-thermal regime onfloweringin cowpea accession TVu 1188. Response surfaces
show (a) effects of photoperiod and night temperature on days from sowing to flowering (/) and
(b) effects of photoperiod and mean diurnal temperature on days from sowing tofloweringif and
l//on right and left hand ordinates, respectively). Solid circles denote the experimental mean values
in each of 12 environments. The linep0 indicates the critical photoperiod (see text), which marks the
intersection of the photoperiod and temperature response surfaces.

the experiment was terminated after 95 days, data for the appearance of ripe pods are
incomplete. The values of the temperature constants of the regressions drawn in Figs 1
and 4 are given in Table 3.

Consequences of constraining temperature responses to a common base temperature

Base temperatures for successive stages of development were calculated for each
genotype (Tables 2 and 3) using Eqn (2) and substituting days from sowing tofirstvisible
bud and first ripe pod for the number of days from sowing to the onset offloweringin
Eqn. 1. Durations to first flower were probably subject to less error than the other two
stages of reproductive ontogeny (both of which involved slightly more subjective
decisions, according to the criteria used here, than 'corolla colour visible'), and the
calculated values of the temperature constants for photoperiod-insensitive accessions are
probably more accurate than those for the photoperiod-sensitive ones (they were
calculated from larger numbers of data points since all stages were achieved within the
duration of each experiment). Base temperatures calculated for time to flowering of
photoperiod-insensitive genotypes varied only from 81 to 10-8 C (Table 2) and we
speculated that all genotypes may, in fact, have a common base temperature or, at least,
that the error introduced by this assumption would be small. This possibility was tested
by calculating regressions for successive stages of development of all genotypes but
constraining the regression lines to converge at the same base temperature. When this
was done, the error was found to be minimized when the base temperature was fixed
at 8 C. Furthermore, this constraint had only a nominal influence on the accuracy of
predicted durations of any stage of development for any genotype. Differences compared
with individual, non-constrained regression predictions were usually less than 2 days (Figs
1,2 and 4). Accordingly, we conclude that for most situations a base temperature of 8 C
is appropriate for cowpeas. Only one accession (TVu 3699) out of the 11 tested seems
to respond in a manner appreciably different to the general theme (and then only with
540 Hadley et al.Reproductive Development in Cowpea

TABLE 3. Values of constants in Eqn (1) derived from regressions of respective inverse
durations {daysfrom sowing)for first visible buds, first openflowersand first ripe pods against
mean diurnal temperature for each of five photoperiod-sensitive genotypes of cowpea

Standard Base
error temperature
Genotype Intercept (a) Slope (b) of slope CO (-a/*) r*

First visible buds

TVu3 -00113 000184 000026 61 0-587
TVu 167 -0-0113 0001 70 0-00029 6-6 0-680
TVu 1047 -00200 000229 000022 8-7 0-809
TVu 1188 -00090 000110 000011 8-2 0-883

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TVu 3699 -00598 000378 000016 15-8 0-971
First open flowers
TVu 3 -00125 000145 000013 8-6 0-772
TVu 167 -00120 000139 000016 8-6 0-818
TVu 1047 -00197 000178 000010 110 0-921
TVu 1188 -00069 000129 0-00005 5-3 0-979
TVu 3699 -00362 000253 000014 14-3 0-944
First ripe pods
TVu 3 -00086 000095 000006 91 0-894
TVu 167 -00068 000090 000009 7-2 0-873
TVu 1047 -00126 000210 000011 60 0-862
TVu 1188 -00023 000075 000004 31 0-969
TVu 3699 -00234 000155 000010 151 0-960

respect to the value of the base temperature) and we speculate in the Discussion as to
why the base temperature for this 'weedy' genotype (IITA, 1974) should be much greater
(Table 3) than the value applicable to other cultivars.
Assuming that a base temperature of 8 C is appropriate for cultivated cowpeas, the
thermal sums required to reach particular stages of reproductive development in any
genotype in all environments can now be calculated relative to a common baseline. The
thermal sums presented in Table 4 can be used to interpret the progress towards
reproductive maturity of cowpea cultivars grown in fluctuating field environments, and
especially of those genotypes insensitive to photoperiod, providing, of course, that
adequate meteorological data are available. Furthermore, applying a common base
temperature to all genotypes simplifies considerably the task of developing screening
techniques for responsiveness to photo-thermal conditions, as we discuss below.

DISCUSSION
Previous work on photoperiod-temperature effects on reproductive development in
cowpeas and other grain legumes (e.g. Huxley and Summerfield, 1974) has emphasized
the importance of night temperature per se. Whilst this emphasis seemed justified from
the data derived from small factorial experiments, and was supported by observations
in thefield(Littleton et al., 1979). It is now apparent that genotypes are sensitive to mean
diurnal temperature rather than to night temperature.
This present work has shown that the rate of reproductive development in cowpeas
is a simple function of either photoperiod or mean diurnal temperature over wide ranges
of each of these factors. When photoperiod has no effects on development (i.e. in any
photoperiod for insensitive genotypes or in inductive photoperiods for the sensitive ones),
times to first visible bud, first flower and first ripe pod all show an inverse linear
relationship with mean diurnal temperature. In photoperiod-sensitive genotypes, the
 Hadley et al.Reproductive Development in Cowpea 541
T A B L E 4. Thermal sums {day degrees above 8 C)for the appearance, respectively, of visible
buds, open flowers and ripe pods in diverse genotypes of cowpea*

Thermal sums to the appearance of

Genotype Visible buds Open flowers Ripe pods

Photoperiod-insensitive
TVu 176 377 (2) 621 (2) 997(3)
TVu317 505(9) 751(10) 1127(10)
TVu 465 493(7) 691 (6) 1049 (5)
TVu 946 437 (4) 635 (3) 939(1)
TVu 1009 408(3) 671 (5) 1071 (6)

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TVu 1503 358(1) 594(1) 958 (2)
Mean 430 661 1024
Difference between stages 231 363
Photoperiod-sensitive
TVu 3 503(8) 721 (8) 1123(9)
TVu 167 540(10) 748(9) 1076 (7)
TVu 1047 460(6) 704(7) 1096 (8)
TVu 1188 447(5) 661(4) 1008(4)
Mean 488 709 1076
Difference between stages 221 367
TVu 3669t 553 810

* Calculated when regressions (Eqn 1) are constrained to a common base temperature of 8 C. Figures in
parentheses rank the genotypes in order of earliness based on thermal sum.
t Atypical response, thermal sums subject to large errors as calculated here (see text).
Dash denotes that ripe pods were not formed within the duration of the experiment.

delay in flowering increases inversely with increasing photoperiod beyond a critical value
which varies with temperature.
One objective of this research programme has been to develop techniques for screening
large numbers of genotypes which may be more or less sensitive to photoperiod and
temperature (and which, therefore, are likely to be well adapted in either a relatively few
or a wide range of environments, respectively). The technique developed and used already
(Summerfield, Minchin and Wien, 1979) can now be improved in the light of the results
discussed here.
The concept of a temperature-time integral above a base temperature, variously
described as 'day-degrees', 'heat units' or 'thermal time', has often been used to predict
developmental processes in plants. It has not always been realized, however, that the
concept is only valid if a function of the type exemplified by Eqn (1) applies (Monteith,
1977; Angus et al., 1980). The concept has been used recently to describe the effects of
temperature on rate of germination (Garcia-Huidobro, Monteith and Squire, 1982a, b),
and the evidence provided here reinforces the value of this approach for reproductive
development too. The special value of the 'thermal time' concept is that it can be applied
to fluctuating temperature environments and, where the base temperature is known, it
would be possible to calculate the temperature response surface from the results obtained
in a singlefieldenvironment. For example, it has been shown here that for many cultivars
little error is introduced by assuming a common base temperature of 8 C. Thus, by
recording the tjrhe taken for cowpeas toflowerin a single location, the value of b could
be calculated/from Eqn (3), then the value of a from Eqn (2), and finally the entire
response surface by inserting these values into Eqn (1).
Similar estimates could be made for photoperiod-sensitive cultivars too, providing that
they are evaluated in inductive photoperiods (say 10 h), and controlled daylength
542 Hadley et al.Reproductive Development in Cowpea
glasshouses would seem admirably suitable for this purpose (Summerfield et al., 1979).
Even if such a facility were not available, we suggest that a prudently-selected field site
and growing season (days shorter than 12 h and mean temperatures close to 20 C) might
prove an acceptable and reliable alternative. The thermal times calculated (as in
Table 4) could then be used to predict reproductive development in other fluctuating
field environments where adequate meteorological data were available.
However, although a base temperature of 8 C is apparently typical, accession TVu
3699 was an exception. This is a 'weedy' type first collected from the hot, semi-arid,
sub-Sahelian zone of northern Nigeria (IITA, 1974) where typical mean maximum
temperatures are close to 35 C during thefirstfew weeks of the rainy season and usually
exceed 30 C thereafter (Wien and Summerfield, 1980). When the regressions for this
genotype were constrained to 8 C large errors were introduced and predictions were not

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reliable. Independent regressions indicate that it has a base temperature of 15 to 16 C
(Table 3). We suggest that the acute sensitivity to photoperiod shown by this genotype
(Fig. 1; Lush et al., 1980) is 'reinforced' by a warmer than usual base temperature for
reproductive development. These two responses combined would ensure timely develop-
ment in the indigenous climate so that plants come into flower at the end of the rainy
season and then produce their pods when climatic conditions are less likely to cause
spoilage. The existence of such a genotype therefore suggests that a temperature screen
should, ideally, be based on at least two thermal environments in order to expose such
anomalies.
It is clear from Fig. 5(b) that a screen for photoperiod sensitivity would be most
reliable if carried out in a warm mean temperature (about 28 C) so that temperature
effects did not interfere with those due to photoperiod. Because the reciprocal of time
to flowering is linearly related to photoperiod, two environments with different
daylengths would be sufficient to estimate the values of the photoperiod constants a' and
b' in Eqn (4). Natural, short tropical days can be extended easily on a field scale using
incandescent lamps suspended above the crop (Summerfield et al., 1979). Thus, from two
simple screens, it should be possible to estimate the entire photo-thermal response surface
for any genotype from two planes (determined from Eqns 1 and 4) which intersect at
the pc given by Eqn (5) and shown in Fig. 5(b).
We believe that the model based on Eqns (1) and (4) describes the principal features
of the photo-thermal regulation of reproductive development in cultivated cowpeas.
Furthermore, we provide evidence elsewhere (Hadley, Summerfield and Roberts, 1982)
that Eqn (1) applies to other grain legumes and that, with some modifications, similar
types of photoperiod-temperature models may also be applied, in particular, to Glycine
max, Phaseolus vulgaris, Pisum sativum and Vigna radiata. In the present investigations,
the 12 different thermal regimes were created by varying night temperature and the
periods in each diurnal cycle for which day and night temperature were imposed.
However, in the other grain legumes, evidence from regimes created by variations in day
temperature was also included. We also discuss elsewhere (Hadley et al., 1982) limitations
to these relationships which might occur under conditions of thermal stress, created by
excessively hot days or cold nights.
Whereas it is clear that the form of the temperature relationship shown here lends itself
to treating a fluctuating temperature environment in terms of thermal units, we would
not wish to imply, by analogy, a similar treatment could be applied in terms of
photoperiodic units. It is clear in cowpea, as in other species, that sensitivity to
photoperiod changes ontogenetically (Summerfield et al., 1982; and see Jones and Laing,
1978 for soyabeans) and, of course, several species have a juvenile phase during which
they are insensitive to photoperiod (Vince-Prue, 1975).
The model described here does not explain the underlying mechanisms which control
reproductive ontogeny, but it defines what now has to be explained. Furthermore, we
 Hadley et al.Reproductive Development in Cowpea 543
believe that it enables, for the first time, adequate design of simple experiments to
investigate the separate genetic control of photoperiodic and temperature responses. Such
experiments might help to explain the phenotypic interactions between the genes which
control reproductive ontogeny in these sensitive species.

ACKNOWLEDGEMENTS

This work, in collaboration with the International Institute of Tropical Agriculture,

Nigeria, was generously funded by the UK Overseas Development Administration. The
engineering assistance of Messrs D. Dickinson, A. C. Richardson, K. Olivers, S. Gill
and C. Billing and technical assistance of Mrs B. Whitlock, Miss C. Chadwick, Mrs K.
Knott, Messrs T. Pitman, M. Craig, M. Boxall and J. Franklin is gratefully acknowledged.

Downloaded from http://aob.oxfordjournals.org/ at National Sun Yat-sen University on November 6, 2016

The Root Nodule Pty Ltd., Epping, NSW, Australia kindly provided the Rhizobium.

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