Expl Agric. (1990), volume 26, pp.

31-40
Printed in Great Britain

AN AGRONOMIC EVALUATION OF JACKBEAN (CANAVALIA

ENSIFORMIS) IN YUCATAN, MEXICO. III. GERMPLASM
ByC. D.J.KESSLERf
Facultad de Medicina Veterinaria y Zootecnia, Universidad Autonoma de
Yucatan, AP 116-D Merida, Yucatan, Mexico 97100 and School of Plant
Biology, University College of North Wales, Deiniol Road,
Bangor, Gwynedd LL57 2UW, Wales
(Accepted 12 December 1988)

SUMMARY
A field experiment was carried out to compare the performance of 18 accessions of jackbean
from different origins. Plant development and phenology, final plant biomass, and seed yield
and its components were evaluated. Flowering time ranged from 56 to 109 days after sowing.
Bush and climbing types were identified and differences in seed yield were recorded, the largest
yield occurring in a mid-season flowering accession. It was concluded that more material should
be screened since germplasm introduction appears to be a promising route to larger jackbean
yields in Yucatan.

C. D. J. Kessler: Evaluation agrondmica de canavalia ensiformis en Yucatan, Mexico. III.

Germoplasma.

RESUMEN
Un experimento de campo fue llevado a cabo para comparar el comportamiento de 18 accesiones
de C. ensiformis provinientes de ori'genes distintos. El desarollo, la fenologi'a y la biomasa final
de las plantas, asf como el rendimiento de la semilla y los componentes del rendimiento de la
semilla fueron evaluados. El tiempo de la siembra hasta la floracion tuvo un rango de 56 a 109
dias. Se identificaron tipos arbustivos y tipos enredaderas y se registraron variaciones en el rendi-
miento de semilla, el mayor encontrandose en una accesion de floracion intermedia. Se con-
cluyo que mas material debe ser probado ya que la introduction de germoplasma parecio ser
una ruta prometadora a rendimientos mayores de la C. ensiformis en Yucatan.

INTRODUCTION

Grain legumes generally exhibit a range of biological diversity. For example in

common bean (Phaseolus vulgaris), a widespread grain legume, genetic variation
in seed size, growth habit, disease and pest resistance, adaptation to photo-
period and temperature, tolerance of poor soil and drought, and adaptation to
intercropping has been described (Adams et al., 1985). Similar variation in
cowpea (Vigna unguiculata) and soyabean (Glycine max) has also been des-
cribed (Steele et al., 1985; Hymowitz and Newell, 1980). On the other hand in
jackbean, it has been claimed that very little diversity exists (NAS, 1979), and
the agronomic studies on jackbean reported by Kessler (1990a, b) employed
just one accession, of Venezuelan origin. This paper describes a preliminary
f Present address: Commission of the European Communities, Rue de la Loi, 200, B-1049 Brussels,
Belgium.
32 C. D.J.KESSLER

field evaluation of 17 other jackbean accessions of differing origins, and gives

particular emphasis to interpreting differences in final seed yields in morpho-
logical and phenological terms. Flowering time was also of special interest, par-
ticularly because early flowering should allow adaptation to seasons when a
short duration of rainfall leads to reduced seed yield (Kessler, 1990a, b).

MATERIALS AND METHODS

The experiment was carried out in Yucatan, Mexico, on land cleared of secon-
dary bush. The soil was red, derived from limestone, of pH 7.8-7.9 and about
40 cm deep. General site and weather conditions, crop husbandry and evalua-
tion techniques have been described in a previous paper (Kessler, 1990a). The
accessions and their origins are listed in Table 1. PI material was received from
the United States Department of Agriculture, Regional Plant Introduction
Station, Georgia, USA, and accessions will be identified in the text by their
first three digits only; H material was received from Universidad Central de
Venezuela, Maracay, Venezuela. Seeds were sown on 3 July 1985 in four
blocks. Plots were square and consisted of nine plants at a density of 2.25
plants m~2 surrounded by guard rows of the original jackbean material which
was pruned where necessary to the form of the plants in the plot.
Accessions were categorized into bush or climbing types, and according to
the time to flowering. Bush-type plants were compact, erect, self-supporting,
formed a canopy at about 80-100 cm above ground level and developed stiff
lateral branches at the lower nodes. Climbing-type plants were lax, tended to
wind their stems around supports and had internodes that were about twice the
length of those of the bush types; branches, which arose at the lower nodes,
and the main stems were often over 2 m in length. Accessions that displayed
the climbing habit were supported on an open horizontal framework of parallel
wooden bars placed at a height of 80 cm.
The height of all plants was measured 42 days after sowing (DAS), and the
number of nodes and branches counted. Three accessions were selected at
58 DAS from those which flowered first and, in each plot, three plants were
chosen at random for height measurement and a count of the number of nodes
on the main stem. At 111 DAS, the percentage ground cover of all accessions
Table 1. Jackbean accessions and countries of origin
Accession Origin Accession Origin

PI181048 India PI364355 Mozambique

PI192975 Guadeloupe PI404610 Paraguay
PI276655 St Kitts PI451828 Guatemala
PI279593 Philippines H2a Venezuela
PI284789 Sudan H2b Venezuela
PI308540 Colombia H5 Venezuela
PI311504 Brazil H7 Venezuela
PI338584 Costa Rica H12 Venezuela
PI358592 Ethiopia Original Venezuela, original 1980
introduction to Yucatan
 Jackbean in Yucatan, Mexico. Ill 33

was estimated by eye and in the early-flowering accessions, the number of

nodes on the main stem was counted and plant height and diameter (mean of
two measurements taken at right-angles) were measured.
When pods were mature, seed yield and its components were recorded in all
accessions and the biomass of above-ground plant parts in seven accessions
chosen to represent different plant form and flowering-time categories. Seed
was harvested from all the plants in a plot, and biomass recorded on two repre-
sentative plants. Samples were dried at 60C to a constant weight. In addition,
the length of the main stem and branches, the number of nodes on the main
stem, the node positions of branches and peduncles on the main stem and of
pods on peduncles, and the number of leaves and leaf scars were recorded.
Secondary and the occasional higher order branches were classed together and
branches less than 5 cm long were ignored. Harvest index was calculated as the
weight of seed as a percentage of total above-ground plant weight.
In the early flowering accessions whose biomass was not analysed, two
representative plants in each plot in three blocks were selected at the time of
the seed harvest for recording the height and number of nodes on the main
stem, the number of nodes with branches and branch length, and the number
of leaves and leaf scars. Branches less than 5 cm long were not included.

RESULTS

Early vegetative growth

A week after sowing all accessions except PI404 had emerged uniformly.
PI404 was not evaluated since a large proportion of the seeds appeared to be
hard and took up to six months to germinate. The early growth of all accessions
appeared similar; however, PI451 was notable for having very dark green foliage
and a large leaflet size. PI279 also had dark green foliage, and stems which were
reddish-coloured from an early age: normally stems were green and became
reddish only at maturity.
By 42 DAS, there were some clear differences in the pattern of vegetative
development among the accessions, which were grouped according to flowering
time and growth habit (Table 2).

Flowering
There was a well-defined group of accessions which flowered at around 58
DAS and these, together with PI279, are classed as early flowering. PI451
followed at 84 DAS and is classed as mid-season flowering, and the rest of the
accessions, which flowered at 91-109 DAS, are classed as late flowering. Early
and mid-season flowering accessions had a much shorter flowering period than
those which flowered later. All accessions had a single flowering period, during
which flowers appeared on the different racemes in an overlapping sequence.
The last observations of flowering were made at 114 DAS when not all plants
had flowered, but all did eventually flower and set pods.
34 C. D.J.KESSLER

Table 2. Plant form at 42 DAS, mean flowering time and classification

of jackbean accessions
Plant No. nodes No. Days
Flowering Plant height on main branches to 50%
Accession time form (cm) stem > 1.0 cm flowering

PI192 early bush 32 6.9 3.1 56

PI181 38 7.4 3.8 57
PI308 35 7.0 3.3 57
PI358 i "n 33 6.3 3.0 57
PI311 1 t 34 7.3 3.5 58
PI338 35 6.9 2.8 58
PI284 1 l 34 6.8 2.8 60
PI 364 , 31 6.5 2.4 60
PI276 t >> 28 6.0 2.4 62
PI279 ft 39 7.6 4.1 65
PI451 mid-season ,, 43 6.4 2.0 84
H12 late 46 6.8 1.4 91
Original 40 6.0 1.0 93
H5 climbing 68 6.0 1.2 95
H2b it 67 6.0 1.3 100
H2a 11 a 91 7.4 1.6 101
H7 > 73 6.3 0.9 109
LSD (P = 0.05) 12 1.0 1.4 5

Vegetative growth after flowering

The development of the nodes on the main stem and the height of three
early flowering accessions, PI284, PI181 and PI192, followed a similar pattern;
at the time of flowering, 58 DAS, they had a mean height of 58 cm and node
number of 11.3. However, when flowering had finished, at 111 DAS, the mean
height had increased to 77 cm and the number of nodes to 18.0, and these
values, which were similar in all early flowering accessions, did not then alter
until pod harvest. Ground cover at 111 DAS was 60-80% in the early flowering
accessions and almost 100% in all the others.

Plant structure at seed harvest

Seed harvests were made on 161, 191 and 225 DAS in the early, mid-season
and late flowering accessions, respectively, when pods were ripe and most of
the plants had dried off and shed their leaves.
Diagrammatic representations of the branching pattern and pod distribution
of representatives of the four jackbean flowering time and growth habit classes
at the time of pod maturity are given in Fig. 1. The number of nodes on the
main stem of plants in all classes was similar, possibly because the tops of the
main stems of the mid-season and late flowering types tended to dry off earlier
than the rest of the plant. There were large differences in plant form and
branching patterns between the categories. In the early flowering accessions,
plants were small, reaching 77 cm in height and short, single branches were
evenly distributed on the main stem. In the sequence from mid-season flower-
ing to late flowering bush and climbing types, plant height was 140, 128 and
 Jackbean in Yucatan, Mexico. Ill 35

21 (a) Early Mid-season Late climber

19 bush bush
17

12

T
11 1
10
9

44
8
7
6
5
4
3
E
2
1 F
J
n=i
IE
20 0 20150 50 0 50 150150 50 0 50 150250 150 50 0 50 150
Branch length (cm)

1 0 1 2 1 0 1 2 3 2 1 0 1 2 3 2 10 12 3
Branch number
8 " (W ft
7
6
n_
% 5[ |_
i 4 i
3 i
2 [~
1
J
30 20 10 0 20 10 0 10 20 2010 0 10 20 2010 0 10 20
Distribution of pods (%)
Fig. 1. (a) Branching patterns and (b) percentage distribution of pods in jackbean accessions. The vertical
axis represents the main stem starting at the base of the plant, J representing the juvenile leaf node. The
open and shaded histograms represent the total length of branch or the distribution of pods at each node
on the main stem or primary branches, respectively. Bars represent the number of branches in each
category. Early bush refers to PI308 and PI311, mid-season bush to PI451, late bush to the original
accession and late climber to H5, H2a, and H7. N.B. the larger scale of branch length given to the early
bush accessions.

200 cm, respectively, total branch length increased more than 10-fold and
became more concentrated at the lower nodes, and secondary branches increased
both in number and in length. The difference in the length of main stem and
branches between the bush and the climbing accessions was due to the length
of the internodes, since the number of nodes on the main stem was similar in
the two types. The forms of plants in all the early flowering accessions, includ-
ing PI308 and PI311 which were examined in more detail, were broadly similar.
Pods were normally borne singly on the peduncles and although up to three
pods were recorded on some peduncles, others carried no pods. The peduncles
arose either at nodes on the main stem or on branches, and in the later-flowering
and climbing types, pods were concentrated lower on the plants and on branches.
36 C. D.J.KESSLER

60 Early bush
50
40
30
20
g 10
to 0

1 30 Mid-season bush
| 20
Z. 10
c 0
o
3 40 Late bush
= 30
| 20
10
0
30 Late climber
20
10
0
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16
Peduncle position
Fig. 2. Percentage distribution of ripe pods on individual jackbean peduncles. The horizontal axis repre-
sents the flowering positions on the peduncle, of which Position 1 is basal or lowermost; genotype group-
ings as in Fig. 1.

The positions of pods on the peduncles are shown in Fig. 2. There was a general
trend from a concentration of pods at the lowest peduncle positions in the
early flowering types to a distribution in which the pods were more evenly
spaced along the peduncle in the late flowering climbing accessions.

Seed yields and components of yield

Seed yields (Table 3) of all the early flowering accessions were similar, with
a mean of 2150 kg ha"1; however, the mid-season flowering PI451 had a much
higher yield, and that of H7, the last accession to flower, was much lower. Seed
yield was positively correlated (r2 = 0.81) with the number of pods per plant
but appeared to be unrelated to single seed weight and seed number per pod. It
was also correlated with plant biomass (r2 = 0.90), with an overall harvest index
of 44%, although this tended to be lower in the later-flowering accessions. Plant
biomass, on the other hand, was similar in both early and late flowering acces-
sions but was substantially higher in the mid-season flowering PI451. The bio-
mass of the early flowering accessions was concentrated in reproductive struc-
tures, but as accessions flowered later, the proportion of biomass as branch
increased while that as main stem varied little between accessions.
Seeds were white in all the bush-type accessions; however, in the climbing
accessions, seeds were blotched reddish or chocolate-coloured on a brown or
white background. Pod walls were all cream-coloured except in H2a and H7
where they were greyish.
 Jackbean in Yucatan, Mexico. Ill 37

Table 3. Seed yields, components of seed yield and distribution of dry matter
in plants at seed harvest in jackbean accessions
Single Total Dry matter distribution (%)
Seed Pod Seed seed plant
Access- yield number number weight biomass Harvest pod ped- main
sion (kg ha"1) plant"1 pod"1 (g) (g) index wall uncle stem branch

Early flowering bush types

PI192 2025 5.8 11.3 1.39
PI181 2655 6.9 10.5 1.62
PI308 2318 6.4 11.4 1.41 214 49 38 2 7 2
PI358 1913 5.2 11.3 1.44
PI311 2363 6.4 11.5 1.44 222 50 39 2 6 2
PI338 2093 6.6 9.6 1.46
PI284 2340 6.4 10.2 1.59
PI364 1890 5.7 10.7 1.39
PI276 1733 4.8 10.7 1.48
PI279 2160 6.4 11.8 1.27

Mid-season flowering bush type

PI451 3375 9.8 11.5 1.34 411 47 30 4 7 11

Late floweringr bush types

H12 2543 6.4 11.5 1.53
Original 1913 5.2 10.5 1.55 252 42 25 3 10 20

Late flowering climbing types

H5 2070 6.5 10.1 1.41 198 43 25 3 7 23
H2b 2550 7.3 9.2 1.62
H2a 2070 6.5 9.3 1.53 259 43 22 3 6 26
H7 1350 4.4 9.5 1.41 190 37 24 3 7 29
LSD 563 1.5 0.8 0.18 116 6 5 1 3 6
(P = 0.05)
ra (vs seed yield) 0.81 0.11 0.12 0.90

After the pods matured some new vegetative shoots appeared at the basal
and/or apical nodes of the main stems, especially in the early-flowering acces-
sions. However, these shoots showed limited development and although occa-
sional flowers were also produced, pod formation from them was negligible.

DISCUSSION

The wide range of variation shown by jackbeans in agronomically important

characteristics such as flowering time, growth habit and seed yield is similar to
that found in other grain legumes such as common bean or cowpea (Adams et
al, 1985; Steele et al., 1985). However, there was little variation among the
early flowering bush-type accessions, which might account for the general lack
of variation reported in the species by NAS (1979).
From the immediate practical point of view, the most important result was
the high seed yield of PI451, although this may simply have arisen because the
accession was better suited to the calcareous soil of the experimental site. The
38 C.D.J.KESSLER

economic legumes frequently show genetic adaptation to concentrations of

major and minor nutrients and to the common toxic elements (Adams and
Pipoly, 1980), and it is relevant to note that PI451 originated in neighbouring
Guatemala where calcareous soils also exist. Alternatively, its high seed yield
might have arisen because it flowered late enough to allow a long crop duration,
but early enough for seed yield formation to avoid the dry season.
The importance of the climbing habit in determining productivity was diffi-
cult to assess because it was associated with late flowering. The climbing acces-
sions tended to have reduced harvest indices, possibly as a result of water stress
during seed-filling (Fischer and Turner, 1978) or because of increased branching.
But pods of the late-flowering accessions tended to be spread out along the
peduncle, suggesting the possible abortion of some reproductive structures.
In grain legumes such as cowpea and common bean, most pods are set from
the first flowers to open (Summerfield et al., 1985; Laing et al., 1984), and this
pattern was also evident in the early flowering jackbean accessions. However,
abscission of buds before flowering has been observed in common bean grown
in unfavourable photoperiods (Ojehomon et al., 1968), and in cowpea in dense
plantings (Summerfield et al., 1985) and under conditions of drought (Hall
and Grantz, 1981).
From the information available it is not possible to relate the performance of
these jackbean accessions in Yucatan to their country of origin but the late-
flowering types, both bush and climbing, came from Venezuela. The blotched
seed of some of the accessions from Venezuela resembles seed found at archaeo-
logical sites in Arizona (Sauer and Kaplan, 1969) and D. Mann (personal com-
munication) has suggested that the accessions identified here as climbing types
do not belong to Canavalia ensiformis (L.) DC. but to C. plagiosperma Piper.
Whatever their taxonomic status, some of the accessions with the climbing
habit produced large seed yields and may therefore have potential.
Since this experiment has established variation in jackbean, selection objec-
tives may now be considered. The variation in time of flowering in the acces-
sions may be exploited in order to select plants with an optimum time for
flowering, though variability in the length of the growing season in Yucatan
complicates this approach. In a year with an early dry season, an early flower-
ing type may be advantageous; however, the early flowering types, though
indeterminate, showed only limited growth once pods had formed and would
not therefore be able to exploit the growing conditions fully if the growing
season were prolonged. On the other hand, the late flowering types, though
well-suited to a long season, would probably not perform well in a short season.
A similar problem has been noted in cowpea, where it was not considered pos-
sible to develop varieties that could both flower early and respond to later
rains, and varietal intercropping has been proposed as a solution (Hall and
Patel, 1985).
Seed yields of the early-flowering accessions might have been larger if the
plant population had been greater but this is not practical for jackbean in
 Jackbean in Yucatan, Mexico. Ill 39

Yucatan because of the increased labour requirement and the rocky soil. Thus
in contrast to the unbranched crop ideotype sought in many species (Donald
and Hamblin, 1983), jackbean for Yucatan should perhaps have a branched
habit, capable of maximizing light interception even when sown at low density.
Such a plant type may have a low harvest index, though this remained remark-
ably constant in experiments with the original accession of jackbean despite
large differences in plant biomass and branching (Kessler, 1987). The harvest
index recorded in jackbean was at the low end of the range recorded in grain
legumes (Laing et al., 1984) but may respond to selection. Due to the mosaic
of rocky ridges and red-soil-filled hollows that characterizes the terrain of
Yucatan, it may be appropriate to sow a jackbean type that can respond to
high plant density on the red soil, which is more uniform and in which sowing
is easier, and to sow the more spreading type on the rocky soil where sowing is
more difficult.
The climbing habit in jackbean may be advantageous for intercropping with
maize, although the maize is more likely to be affected by a climbing than by
a bush type of jackbean; however, the late flowering associated with the climb-
ing habit, though increasing complementarity between the species, will probably
be a disadvantage in all but the longest growing seasons.
This experiment has demonstrated variation in agronomically important
characteristics such as seed yield, flowering time and growth habit among
jackbean accessions. The work should be continued in other seasons and an
attempt should be made to screen more material since the experiment shows
germplasm introduction to be a promising avenue to higher jackbean yields in
Yucatan.

Acknowledgements. This work was carried out whilst the author was a Techni-
cal Cooperation Officer of the Overseas Development Administration (UK
Government) and the field work was partially supported by the International
Foundation for Science (Sweden). The jackbean accessions were supplied by
Dr A. Escobar, Universidad Central de Venezuela, Maracay, Venezuela (H
accessions) and by United States Department of Agriculture, Regional Plant
Introduction Station, Experiment, Georgia, USA through the good offices of
Dr G. A. White (PI accessions). Finally, I am particularly grateful to Dr C.
Marshall for help with preparation of the manuscript.

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