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University of Notre Dame Australia
[email protected]
Sciences Papers and Journal Articles
School of Sciences
2011
Growth, yield and seed composition of native Australian legumes with potential as
grain crops
Lindsay W. Bell
CSIRO, [email protected]
Megan H. Ryan
UWA
Richard G. Bennett
UWA
Margaret T. Collins
Centre for Legumes in Medeiterranean Agriculture
Heather J. Clarke
UWA and University of Notre Dame Australia, [email protected]
Follow this and additional works at: http://researchonline.nd.edu.au/sci__article
Part of the Physical Sciences and Mathematics Commons
This article was originally published as:
Bell, L. W., Ryan, M. H., Bennett, R. G., Collins, M. T., & Clarke, H. J. (2011). Growth, yield and seed composition of native Australian
legumes with potential as grain crops. Journal of the Science of Food and Agriculture, 92 (7), 1354–1361.
http://doi.org/10.1002/jsfa.4706
This article is posted on [email protected] at
http://researchonline.nd.edu.au/sci__article/43. For more information,
please contact [email protected]
Research Article
Received: 17 May 2011
Revised: 21 July 2011
Accepted: 16 September 2011
Published online in Wiley Online Library:
(wileyonlinelibrary.com) DOI 10.1002/jsfa.4706
Growth, yield and seed composition of native
Australian legumes with potential as grain
crops
Lindsay W Bell,a∗ Megan H Ryan,b Richard G Bennett,b,c Margaret T Collinsd
and Heather J Clarked†
Abstract
BACKGROUND: Many Australian native legumes grow in arid and nutrient-poor environments. Yet few Australian herbaceous
legumes have been investigated for domestication potential. This study compared growth and reproductive traits, grain yield
and seed composition of 17 native Australian legumes with three commercial grain legumes.
RESULTS: Seed yields of seven native legumes were >40% of Cicer arietnum, with highest seed yields and harvest indices in
Glycine sp. (14.4 g per plant, 0.54 g g−1 ) and Lotus cruentus (10.2 g per plant, 0.65 g g−1 ). Five native species flowered earlier
than field pea (Pisum sativa) (109 days), though many were slower to flower and set seed. Largest seeds were found in Glycine
canescens (17 mg), with seed of other native species 14 times smaller than commercial cultivars. Seed composition of many
native legumes was similar to commercial cultivars (200–330 g protein kg−1 dry weight (DW), 130–430 g dietary fibre kg−1
DW). Two Cullen species had high fat content (>110 g kg−1 DW) and Trigonella sauvissima had the highest crude protein content
(370 g kg−1 DW).
CONCLUSION: The seed composition and reproductive traits of some wild native Australian legumes suggest they could offer
potential as grain crops for soils and environments where the current grain legumes are uneconomic. Further evaluation of
genetic diversity, especially for seed size, overall productivity, and reproductive development is needed.
c 2011 Society of Chemical Industry
Keywords: novel crops; phenology; perennial; Kennedia; Swainsona; Rhynchosia
INTRODUCTION
The performance of many exotic legumes used in Australian
agriculture is constrained by variable climatic conditions and
infertile soils.1 Many indigenous Australian legumes may be
more productive than exotic species when grown under lowfertility or drought conditions.2 – 5 For example, under low soil P
concentrations the native legumes Kennedia prorepens F. Muell.
and Lotus australis Andrews produced 32 times and 11 times more
biomass than lucerne (Medicago sativa), respectively.4 Despite
their inherent adaptation to challenging environments, the diverse
legume flora of Australia has received limited assessment for
agricultural potential, and focus to date has been on their potential
as pastures.5 – 10 Few studies investigate Australia’s legumes for
their potential as alternative grain crops. Furthermore, Australia
has native legumes closely related to globally important grain
legume crops such as Trigonella (fenugreek), Vigna (mungbean),
and Glycine, which includes the most widely grown oilseed legume:
soybean (Glycine max L.). As well as the potential to provide a wider
variety of adapted legumes for modern farming systems, Australian
legume germplasm in these genera could provide plant breeders
across the world with a valuable resource for increased adaptation
to water-limited and infertile environments.
Many Australian herbaceous legumes are also perennial and
offer sustainability benefits for agriculture. Ongoing work is
J Sci Food Agric (2011)
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examining approaches to integrate perennial pastures and tree
crops into agricultural systems to reduce dryland salinity. Perennial
grain crops have been proposed as an alternative that brings
many sustainability benefits to agriculture while maintaining grain
production.11,12 Perennial grain crops may only require grain yields
of 40–60% of current crops if they could grow in areas where
current crops are less profitable or if they could provide additional
grazing for livestock.13 Many of Australia’s native legumes have
potential as forage species6 and hence could be used for the dual
purposes of grain and grazing.
∗
Correspondence to: Lindsay W Bell, CSIRO Ecosystem Sciences, PO Box 102,
Toowoomba, QLD 4350, Australia. E-mail: [email protected]
†
University of Notre Dame Australia, Fremantle, WA 6959, Australia.
a CSIRO Ecosystem Sciences, Toowoomba, QLD 4350, Australia
b School of Plant Biology and Institute of Agriculture M081, University of Western
Australia, Crawley, WA 6009, Australia
c CSIRO Ecosystem Sciences, Wembley, WA 6913, Australia
d Centre for Legumes in Mediterranean Agriculture (CLIMA), University of Western
Australia, Crawley, WA 6009, Australia
c 2011 Society of Chemical Industry
www.soci.org
LW Bell et al.
Table 1. The 17 native Australian herbaceous legumes and three commercial grain legumes grown in the study, the cultivar/accession used and a
description of the growth habit and life cycle of each species
Species
Accession no./cultivar
Growth habit
Native legumes
Cullen australasicum (Schltdl.) J. W. Grimes
Cullen cinereum (Lindl.) J. W. Grimes
Cullen graveolens (Domin.) J. W. Grimes
Cullen tenax (Lindl.) J. W. Grimes
Glycine canescens F. J. Herm
Glycine sp.b
Glycyrrhiza acanthocarpa (Lindl.) J. M. Black
Kennedia coccinea Vent.
Kennedia prorepens F. Muell.
Lotus cruentus Court
Rhynchosia minima (L.) DC
Swainsona canescens (Benth.) F. Muell.
Swainsona colutoides F. Muell.
Swainsona kingii F. Muell.
Swainsona purpurea (A. T. Lee) Joy Thomps.
Swainsona swainsonioides (Benth.) J. M. Black
Trigonella sauvissima Lindl.
SA44380
AusTrCF 320112
AusTrCF 320184
AusTrCF 320110
NIND001
NIND004
C2N01GA
NS-26828
NS-30323
NF003
NF013
KIMS003
NIND006
NF002
KIMS004
KIMS005
Fortescue collection
Erect sub-shrub
Erect sub-shrub
Erect
Prostrate to erect
Perennial twinning
Perennial twinning
Semi-prostrate to ascending
Twining
Prostrate to ascending
Prostrate to ascending
Prostrate or twining
Prostrate or erect, spreading
Erect
Prostrate or ascending
Erect to spreading
Spreading ascending
Erect to prostrate or ascending
Commercial grain legumes
Cicer arietnum L. (chickpea)
Pisum sativum L. (field pea)
Lupinus angustifolius L. (narrow-leaf lupin)
Rupali
Kaspa
Mandelup
Spreading ascending
Twining, ascending
Erect
a
b
Life cyclea
A
P
A
P
P
?
P
P
P or A
P
P
A
A or P
A or P
P
A
A
A
A
A
A, annual; P, perennial.
Species is unknown.
Although Australian legumes were consumed by Aboriginals,
there is no evidence of any kind of their domestication. The use
of seeds as a food source is mainly documented for several Acacia
species,14,15 though there is evidence of Swainsona galegafolia
(Andrews) R. Br. (smooth Darling pea) being eaten fresh and
tasting similar to common garden pea.16 There are few published
records of the ethnobotany of many Australian native legumes; the
ephemeral growth of many species or their rarity in the vegetation
may have rendered them an unreliable food source. Antinutritional
compounds or toxins (e.g. alkaloids, furanocoumarins, and
hydrogen cyanide), present in many Australian legumes, may also
lower their palatability or edibility. ‘Dilly bags’ (leaching baskets)
were commonly used by Aborigines to remove toxins by soaking
certain seeds and legumes in running water for hours or days.
Despite known toxins in Australian legumes, many secondary
compounds in Australian plants have pharmaceutical uses, and
within many genera and species there is significant variation in
their presence and activity.17,18
A recent review of the potential of Australian native legumes
systematically evaluated 14 genera for their adaptation to arid
and winter-dominant semi-arid climatic regions, as well as
characteristics such as pod indehiscence, growth habit, grain
size, seed composition, and likely presence of antinutritional
compounds or toxins.17 This review found that a number
of herbaceous species are likely to be adapted to dry and
infertile environments and possess many desirable attributes
for domestication as grain crops. These species include Cullen
tenax (Lindl.) J. W. Grimes, Crotalaria cunninghamii R. Br., Glycine
canescens F. J. Herm., Glycyrrhiza acanthocarpa (Lindl.) J. M.
Black, Kennedia prorepens, Rhynchosia minima (L.) DC., Swainsona
canescens (Benth.) F. Muell., Swainsona colutoides F. Muell.,
wileyonlinelibrary.com/jsfa
and Trigonella suavissima Lindl. One previous study found that
Hardenbergia violacea (Schneev.) Stearn, Crotalaria cunninghamii,
and Kennedia nigricans Lindl. were worthy of further investigation
as grain crops, as they possessed relatively large seeds (38 mg,
38 mg, and 16 mg, respectively) and contained crude protein
content of 210–280 g kg−1 fresh weight.19 However, there is
little information on the grain or seed production potential
and seed composition of many native Australian legumes.
Therefore, this investigation was conducted to compare the
growth characteristics, reproductive development, seed yield, and
seed composition of selected species with those of commercially
grown grain legume crops. We found that several native legumes
had similar reproductive development, reproductive allocation,
and seed composition, but smaller seed size than commercial
grain legume cultivars.
METHODS
Experimental design and management
Seventeen native Australian legumes and three commercial
grain legumes (Table 1) were grown in a glasshouse from mid
May to December 2008 at the University of Western Australia,
Crawley, Australia (31◦ 59 S, 115◦ 53 E). This period was chosen
to coincide with the winter–spring growing season of grain crops
in southern Australia, including the three commercial species.
The 17 native species were prioritised based on information in
the literature,17,20 as well as unpublished evaluations (R Snowball
and S Hughes, private communication). Seed for several highpriority species identified by Bell et al.17 was not available (e.g.
Crotolaria species), so either lower-priority or other representative
c 2011 Society of Chemical Industry
J Sci Food Agric (2011)
Growth, yield and seed composition of Australian legumes
species were evaluated. Seed was obtained from SARDI Genetic
Resource Centre, Urrbrae, SA; Australian Tropical Crop and Forage
Resource Centre, Biloela, QLD; Kimseed International, Osborne
Park, WA; Nindethana Seed Service, Albany, WA; and from previous
collections made throughout the arid and semi-arid regions of
Western Australia. In addition to the benefits for controlling pests
and diseases, information on self-pollination compatibility of these
native species is limited, so plants were grown in the glasshouse in
the absence of pollinators (and without hand pollination) to favour
species with higher self-compatible pollination – a desirable trait
in a grain crop.
Seeds of all taxa were scarified, imbibed on dampened filter
paper and sown the following day. Six seeds of each species were
sown per pot, in 10 replicate 300 mm black standard pots with
drainage holes. Pots were filled with potting mix composed of
5 : 2 : 3 (v/v) fine composted pine bark : coco peat : coarse sand,
with 1 kg m−3 superphosphate, 2 kg m−3 extra-fine limestone,
0.3 kg m−3 potassium sulfate, 0.2 kg m−3 macro mineral trace
elements, 1 kg m−3 ammonium nitrate (Agran 34.0), 2 kg m−3
dolomite (Ca/Mg), and 0.5 kg m−3 ferrous sulfate heptahydrate.
Pots were placed in a glasshouse at ambient temperature and
supplied with adequate water and nutrients for growth. To
minimise water and nutrient limitations on plant growth, pots
were watered every 2 days (or less frequently if pots were
still moist near the surface) and fertilised every 2 weeks with
approximately 100 mL of a solution of Phostrogen fertiliser
(14 : 4.4 : 22.4 N : P : K, g g−1 ) made up at the rate of 1 g L−1 of
water. Seedlings were thinned to three plants per pot (i.e. 42
plants m−2 ) at 7 weeks after sowing (i.e. 20 June). This density was
chosen to match recommended crop density for the commercial
grain legume species grown under field conditions (i.e. 30–50
plants m−2 ). Because all species grown are indeterminant, water
was withheld once the majority of pods had reached maturity:
from 26 weeks after sowing for commercial grain legume species
(i.e. 10 November) and from 31 weeks after sowing for native
legumes species (i.e. 15 December 2008), because of slower onset
of flowering and maturity. In the glasshouse, temperatures were
optimal for plant growth (maxima of 20–30 ◦ C and minima of
5–15 ◦ C) during the vegetative growth period (mid May to late
August), and increased by up to 5 ◦ C after this time (Fig. 1).
Plant growth and seed composition
Over the experimental period, observations of plant growth and
reproductive development were made every 2–3 days, including
days to emergence, days to appearance of first flower, duration
of flowering and podding, pod dehiscence (shattering) rating,
number of primary and secondary branches at harvest, and height
Glasshouse temperature (°C)
35
30
25
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at harvest. Plants were harvested 3 weeks after water was withheld
and separated into pods, seeds, and shoot material. Biomass
samples were dried at 70 ◦ C for 72 h and seeds were air dried
and stored at 25 ◦ C prior to compositional analysis. Some species
did not flower by the end of the experimental period and hence
only vegetative shoot biomass was measured. Seeds were ground
and then analysed by George Weston Technologies (Enfield, NSW,
Australia) for fat content using fat Soxhlet extraction, crude protein
by combustion method, and dietary fibre by phosphate buffer
digestion, following the methods described in the Official Methods
of Analysis of AOAC International.21
Statistical analysis
Data for each variable measured were analysed by general analysis
of variance (ANOVA) in Genstat version 10 (Lawes Agricultural
Trust, Rothamsted Experimental Station, Harpenden, UK, 2007).
The least significant difference values at P = 0.05 are shown
at the base of each table. Principle components analysis using
Genstat was also conducted to investigate correlations amongst
reproductive, growth and seed yield components in 15 of the
species; two species which had missing data were not included.
RESULTS
Plant reproductive development and growth habit
All indigenous legumes flowered later than the commercial cultivars of chickpea and lupin (Table 2). The fastest-flowering native
species were Trigonella sauvissima, Lotus cruentus, Swainsona
kingii, Glycine sp. and Rhynchosia minima (Fig. 2), all which flowered before the field pea cultivar, taking between 74 and 107 days
after sowing or 1219–1763 degree days (Table 2). The longest
species to flower – Glycyrrhiza acanthocarpa – took 202 days or
more than 3000 degree days before the appearance of the first
flower. A number of native species continued to flower until the
final harvest (212 days after sowing). Cullen australasicum, Glycine
sp., Kennedia coccinia, L. cruentus, R. minima, Swainsona colutoides,
S. kingii, S. purpurea, and T. sauvissima had a distinct flowering
period (68–101 days) and completed flowering and pod set by
harvest, though this duration of flowering was significantly longer
than in the commercial cultivars (Table 2).
The plant height at harvest for most native legumes was
similar to chickpea (457 mm), ranging from 245 (S. kingii) to
528 mm (Swainsona swainsonioides); the exceptions were the
climbing–twining Glycine and Kennedia, which were staked and
hence their height at harvest was greater. The degree of branching
varied significantly between the species, with T. sauvissima, Glycine
sp., G. canescens, S. colutoides, and S. swainsonioides exhibiting
apical dominance similar to field pea. Only two native species (R.
minima and K. prorepens) branched more than chickpea, which is
renowned for its highly branched ascending habit. Pod-shattering
scores indicated that four of the native legumes had full seed
retention and five exhibited moderate pod shattering (Table 2).
20
15
10
5
0
11 May
1 Jun
22 Jun
13 Jul
3 Aug
24 Aug 14 Sep
5 Oct
26 Oct
Figure 1. Maximum and minimum glasshouse temperatures during the
time when the native and commercial grain legumes were grown.
J Sci Food Agric (2011)
Seed yield and yield components
Field pea had the highest seed yield and plant biomass of the
legumes grown. Glycine sp. yielded more seed and biomass than
chickpea, while the seed yield of six other native legumes were
50–75% of chickpea and 20–30% of field pea (Table 3). Some
of the native species also had a harvest index similar to that of
the commercial cultivars; in particular, L. cruentus had one of the
lowest total plant biomasses and one of the highest seed yields,
c 2011 Society of Chemical Industry
wileyonlinelibrary.com/jsfa
www.soci.org
LW Bell et al.
Table 2. Comparison of reproductive development, growth habit, and pod-shattering susceptibility of 16 native Australian herbaceous legumes
and three commercial grain legume crops (highlighted in bold) grown in the glasshouse. Species’ means with least significant difference (LSD).
Narrow-leaf lupins became diseased and were not harvested
Species
Cicer arietnum
Lupinus angustifolius
Trigonella sauvissima
Lotus cruentus
Swainsona kingii
Glycine sp.
Rhynchosia minima
Pisum sativum
Swainsona colutoides
Swainsona purpurea
Swainsona swainsonioides
Cullen cinereum
Cullen tenax
Cullen graveolens
Swainsona canescens
Cullen australasicum
Glycine canescens
Kennedia prorepens
Glycyrrhiza acanthocarpa
LSD (P = 0.05)
a
b
c
Days to
first flower
Duration of
flowering (d)
Thermal time
to first flowera
(◦ C d)
Height at
harvestb
(mm)
No. branches
at harvest
Shattering
scalec
60
74
74
81
92
104
107
109
110
111
111
115
116
122
130
132
157
167
202
44
–
101
100
91
79
84
28
83
69
dnf
dnf
dnf
dnf
dnf
68
dnf
dnf
dnf
1003
1219
1219
1333
1509
1707
1763
1797
1815
1833
1833
1909
1927
2038
2189
2230
2741
2967
na
457
–
444
340
245
702
497
1077
438
463
528
363
491
301
425
330
1366
782
444
25.2
–
7.6
19.2
20.0
6.2
28.0
6.8
4.5
13.6
7.6
15.8
19.0
19.0
15.0
19.5
10.6
44.0
14.0
1
1
3
2
?
2
?
1
?
2
?
1
1
1
2
1
2
?
?
9
7
167
230
7.6
n/a
Sum of average daily temperature (i.e. base temperature of 0).
Height of both Glycine species and Kennedia prorepens is of staked plants.
1, full seed retention; 2, moderate (<60% of pods shattered); 3, severe (>60% of pods shattered); ?, not rated; dnf, did not finish flowering
resulting in a harvest index of 0.65. A number of native legumes
(C. australasicum, R. minima, K. prorepens, S. swainsonioides, S.
purpurea, and S. canescens) produced high amounts of plant
biomass, but they had low seed yields and hence very low harvest
indices (<0.10). Notably, we observed these species to have low
pod set in the glasshouse, which may indicate that they require
pollinators to facilitate flower fertilisation.
A clear distinction between the commercial cultivars and
the native legumes was their seed size, with field pea and
chickpea having seeds weighing 260 mg and 189 mg, respectively
(Table 3 and Fig. 2). The native legumes with largest seeds were
G. canescens (17 mg) and R. minima (13.3 mg). Seeds of Swainsona
species were between 2.7 and 5.5 mg, Cullen were around 5 mg
and the two smallest seeded species were T. suavissima and L.
cruentus. Trigonella suavissima and L. cruentus also had the highest
seed number per plant, with a clear inverse relationship between
seed mass and seed number occurring across the range of legumes
tested (Fig. 3).
Principal component analysis showed close correlations between seed mass and seed yield, mainly associated with the
commercial cultivars (Fig. 2). Of the major factors represented in
the two first principal components, time to first flower was inversely correlated with harvest index and seed number per plant,
and biomass and plant height were inversely correlated with seeds
per plant. Amongst the native legumes, the two Glycine species
were differentiated based on both their higher seed yield and seed
mass. In addition to the Glycine species, the six native species in
the top right-hand quadrant were those with the highest seed
yields and harvest index and more rapid onset of flowering (Fig. 2).
wileyonlinelibrary.com/jsfa
Seed composition
Most native legumes had seed composition within the range
of the commercial grain legume cultivars (Table 4). The only
exceptions were R. minima, which had lower crude protein content
(<220 g kg−1 DW) and two Cullen species, with fat content over
110 g kg−1 DW, which was higher than all other legumes tested
here (12–62 g kg−1 DW) (Table 4). Protein content was highest in
T. sauvissima (373 g kg−1 DW), but a number of native legumes
had crude protein greater than 300 g kg−1 . Chickpea and field pea
protein content was lower than most native legumes. Lupin had
the highest dietary fibre content (476 g kg−1 DW), but the native
Glycine and Lotus species also had high dietary fibre (>300 g kg−1
DW). The remainder of the native species had fibre content in the
range 196–286 g kg−1 DW (Table 4).
DISCUSSION
We expected that the seed yield, reproductive allocation and seed
composition of grain legume cultivars would be far superior to the
native Australian legumes, which have received no domestication
or selection for grain yield or quality. However, yields greater than
40% of chickpea and similar reproductive allocation, comparable
phenology, and moderate to low pod shattering were found
in Glycine sp., L. cruentus, S. kingii and S. colutoides. Three
other species – C. cinereum, C. tenax, and G. canescens – flowered
later than the grain legume cultivars (115, 116 and 157 days
after sowing, respectively) but also yielded >40% of chickpea.
Universally, seed size was ∼14 times smaller in the native legumes
than the domesticated species. The highest yields were in species
c 2011 Society of Chemical Industry
J Sci Food Agric (2011)
Growth, yield and seed composition of Australian legumes
www.soci.org
4
PC2
3
2
1
C. graveolens
Kennedia prorepens
TT to flower/DTF
-3
# branches
S. canescens
C. australasicum .
S. swainsonioides
-2
S. purpurea
R. minima
0
-1
Pod #
Trigonella sauvissima
Seed #
Lotus cruentus
C. tenax
S. kingii
S. colutoides
PC1
Seeds per pod
C. cinereum
0
1
2
3
Glycine sp.
Glycine canescens
HI
-1
Chickpea
Biomass yield
Height
-2
Seed yield
Seed mass
-3
Field pea
-4
Figure 2. Principal component analysis of 11 native Australian legumes (filled diamonds) and two commercial grain legume cultivars (unfilled diamonds)
based on plant reproductive development, seed yield, and yield components. Biplot vectors indicate strength and direction of factor loadings for PC1
(y-axis, 29.6% of variation) and PC2 (x-axis, 26.5% of variation).
Log10 seed number plant-1
4.0
y = -0.7538x + 3.1403
R2 = 0.8248
3.0
2.0
L. cruentus
T. suavissima
S. kingii
C. tenax
S. colutoides
Glycine sp.
C. cinereum
G. canescens
C. graveolens C. australasium
R. minima
Field pea
Chickpea
1.0
0.0
0.0
1.0
2.0
3.0
Log10 seed mass (mg)
Figure 3. Relationship between individual seed mass and seed number
produced per plant for native Australian legumes and two commercial grain
legume cultivars. Those native species with poor seed set, presumably due
to pollination problems, are omitted.
that produced a large number of seeds per plant; the one
exception was the largest-seeded native legume: G. canescens.
Many of the seven native legumes with the highest reproductive
allocation and grain yield in this study are also perennials (i.e.
Glycine sp., L. cruentus, C. tenax and G. canescens) and hence offer
J Sci Food Agric (2011)
some potential as perennial grain crops, which could provide
a number of environmental and production benefits in farming
systems.11,12 Overall, we believe that the seven native Australian
legumes mentioned above have adequate growth, yield, and
seed composition to be subject to further investigation for their
agricultural potential as grain crops.
The experiment reported here was conducted under glasshouse
conditions with warmer temperatures, and more favourable
moisture and fertility than would be expected in the field. Yet
the similar plant density, crop productivity, and seed mass in fieldgrown commercial grain legumes provides some basis for these
results to be translated to field conditions. First, plant density
in pots (i.e. 42 plants m−2 ) was in the range recommended for
commercial grain legumes in the field,22 suggesting that interplant competition for light and soil resources was similar to field
conditions. Secondly, harvest index, seed mass, and crop grain and
biomass yield calculated on an area basis are also similar to those
reported for crops grown under favourable field conditions.22
Compared to field-grown crops, chickpea in this study had similar
grain yield (1.9 t ha−1 compared to 1.5–2.0 t ha−1 in the field),
crop biomass (3.3 t ha−1 compared to 3–5.7 t ha−1 in the field),
and seed mass (189 mg per seed compared to 156–186 mg per
seed in the field), though harvest index was higher (0.6 compared
c 2011 Society of Chemical Industry
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LW Bell et al.
Table 3. Seed yield, total plant biomass, harvest index (HI), and yield components of 16 native Australian herbaceous legumes and two commercial
grain legume crops (shown in bold) grown in the glasshouse; three plants were grown in a 300 mm diameter pot. Data are species’ means; least
significant difference (LSD) is provided. Narrow-leaf lupins became diseased and were not harvested
Species
Seed yield
(g per plant)
Total biomass
(g per plant)
HI
(g g−1 )
Seed mass
(mg per seed)
9.94
4.78
4.56
3.40
2.77
2.68
2.18
2.06
2.01
1.43
1.42
0.86
0.82
0.17
0.04
0.02
0.01
n.a
0.94
20.4
8.9
7.7
5.3
8.7
7.8
4.6
6.6
9.6
5.0
15.6
13.4
6.1
10.0
10.6
13.7
13.6
5.2
2.5
0.50
0.54
0.60
0.65
0.30
0.35
0.47
0.30
0.21
0.35
0.09
0.07
0.11
0.02
0.004
0.001
0.001
n.a
0.09
258.9
11.2
188.7
1.5
5.2
16.9
2.7
5.2
3.1
1.2
8.7
13.3
5.7
2.7
3.2
6.3
5.5
n.a
17.7
Pisum sativa
Glycine sp.
Cicer arietnum
Lotus cruentus
Cullen tenax
Glycine canescens
Swainsona kingii
Cullen cinereum
Swainsona colutoides
Trigonella sauvissima
Cullen australasicum
Rhynchosia minima
Cullen graveolens
Swainsona canescens
Swainsona purpurea
Kennedia prorepens
Swainsona swainsonioides
Glycyrrhiza acanthocarpa
LSD (P = 0.05)
Seed no.
per plant
39
428
24
2320
515
159
809
393
645
1185
177
67
141
53
14
1
3
–
210
Pods per
plant
Seeds
per pod
12
79
28
184
515
38
103
393
66
340
177
43
141
17
4
0.6
4
–
120
3.2
5.4
0.9
12.6
1.0
4.2
7.9
1.0
9.8
3.5
1.0
1.5
1.0
3.1
3.5
1.6
0.7
–
2.7
n.a., plants did not produce seed during experimental period.
Table 4. Seed composition of 14 native Australian herbaceous legumes and three commercial grain legume crops (shown in bold) grown in the
glasshouse. Lupinus angustifolius plants grown in the glasshouse and sown at the same time as the native species were diseased; seed was harvested
from some later-sown plants grown in the glasshouse. Owing to insufficient seed, Kennedia coccinea, K. prorepens and Swainsona purpurea were not
analysed
Species
Trigonella sauvissima
Lupinus angustifolius (lupin)
Cullen cinereum
Swainsona kingii
Glycine canescens
Swainsona swainsonioides
Cullen australasicum
Glycine sp.
Cullen tenax
Lotus cruentus
Cullen graveolens
Swainsona colutoides
Swainsona canescens
Pisum sativa (field pea)
Glycyrrhiza acanthocarpa
Cicer arietnum (chickpea)
Rhynchosia minima
Crude protein (g kg−1 DW)
Fat (g kg−1 DW)
Dietary Fibre (g kg−1 DW)
373
369
362
343
340
325
324
322
321
320
304
275
269
263
261
229
210
53
51
118
25
62
36
38
52
113
59
57
21
40
12
n.a.
45
n.a.
204
476
262
236
317
196
286
350
279
429
n.a.
239
227
154
n.a.
257
n.a.
n.a., insufficient seed for analysis.
to 0.4–0.5 in the field). Field pea grown in our experiment seemed
to convert biomass to grain yield more efficiently here than under
field conditions; grain yield was 4.2 t ha−1 compared to 1.5–2.7 t
ha−1 in the field and harvest index was 0.5 compared to 0.35–0.4 in
the field, but seed mass and total crop biomass were similar to field
observations (259 mg per seed compared to 179–240 mg per seed
and 8.5 t DM ha−1 here compared to 6–10 t ha−1 , respectively).22
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These comparisons suggest that under favourable field growing
conditions the best native legumes included in our glasshouse
experiment could yield 1.2–2.0 t ha−1 of grain and 2.4–4.0 t ha−1
of biomass in the field, with similar seed size but possibly reduced
harvest index. Under more arid or infertile growing conditions
these productivity levels would be reduced, but the expected
relative reduction in productivity of native legumes would be
c 2011 Society of Chemical Industry
J Sci Food Agric (2011)
Growth, yield and seed composition of Australian legumes
less than for the exotic legumes.3 – 5 The low seed set in some
species (S. canescens, S. purpurea, S. swainsonoides, K. prorepens,
C. australasicum, and R. minima) is likely to be due to the lack
of pollinators in the glasshouse and confirms the dominance
of open pollination suspected in these species.17 Similarly, the
photoperiod sensitivity of many native legumes is unknown and
the photoperiod regime during the experimental period may have
influenced the growth and development of some species. For
example, long days (>12–13 h) are thought to induce flowering
in C. australasicum,7 which may explain the long period to the start
of flowering observed here. On the other hand, numerous Glycine
species (including soybean) are short-day responsive, suited to
spring sowing, and the high yields of Glycine sown in autumn in
this study suggest that higher yields still might be expected when
sown in spring.
An important consideration for the potential domestication
of native legumes is the end use of the grain, in particular,
concentration of protein, fats/oils, fibre and the presence of
antinutritional or beneficial compounds. In this study we found
that most of the native legumes had higher grain crude protein
content than field pea and chickpea, and some were similar to the
high protein content found in lupin. Previous studies have reported
lower grain protein contents in native Australian legumes,19,23
though it is unclear whether this was calculated on a dry mass
basis. Grain of the two Cullen species had a higher fat content
(>110 g kg−1 DW) than the other native legume seeds tested
here, and fat contents greater than 80 g kg−1 were found in two
species not tested here: K. nigricans and H. violacea.19 Cullen seed
has an adherent pod so fats and oils could be concentrated there,
although, surprisingly, the inclusion of this pod in the analysed
sample did not result in elevated fibre content. Despite the high
oil content in Cullen, these levels are much lower than found
in soybean (∼200 g kg−1 ) and other oilseed crops. A number
of Australian legumes are also known to produce toxins (e.g.
swainsonine in some Swainsona species, hydrogen cyanide in
Australian Lotus) and other bioactive compounds (e.g. isoflavones,
especially phytoestrogens in Glycine, furanocoumarins in Cullen)
which could limit their use in food products. However, a number
of these compounds are pharmaceutically useful and could offer
potential as natural medicines.17 Many cultivated grain legumes
also contain antinutritional compounds that have been lowered by
breeding (e.g. alkaloids in lupins),24 and further analysis to test for
the presence and variation in concentrations of these compounds
within the taxa is required.
In this study we only compared one accession of each species,
yet there could be substantial genetic variation in key agronomic
traits within species, and much more productive genotypes are
likely to exist. This is clearly shown by Snowball et al.,20 who found
that seed yield and plant survival in the field varied considerably
among accessions of native species, including K. prorepens
and S. canescens. Comparisons of seed size and reproductive
development between the present and past studies suggest there
is substantial genetic variation within taxa. For example, in the
present study G. canescens produced two–three times larger seeds
(16.9 mg) than recorded previously for this species (5.9–8.9 mg).17
Others have shown that seed size varies by two–four times within
Kennedia species,17 two times in Glycinelatifolia,25 and greater than
two times in R. minima.26 Drivers of reproductive development are
also variable. Cullen cinereum was much slower to flower in this
study (1900 degree days or 115 days) than previously reported (i.e.
900 degree days or 40 days).27 Flowering in other Cullen species is
thought to be induced by long days and this may also be the case
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in this accession of C. cinereum.7 The T. suavissima accession tested
here flowered more quickly (74 days) than reported previously
for this species (111–118 days),17 suggesting that material that
flowers much earlier may be available. In this study, R. minima
flowered after 107 days, which is midway between that reported
previously (43–142 days).26 Overall, there have been few studies
of variation in agronomic traits of native Australian legumes, but
current evidence suggests there is substantial variation that could
be exploited to improve productivity, seed size, and reproductive
development to fit into different environments.1
CONCLUSION
This preliminary investigation of a range of native Australian
legumes, chosen for this study because of adaptation to
arid and semi-arid winter dominant rainfall conditions, has
shown that, while many species had much lower yields and
reproductive allocation, seven species warrant further evaluation
for domestication for agriculture. These seven species exhibited
grain yields of 40–60% of cultivated grain legumes with grain
protein, fat and fibre in the range desirable in food and
feed industries. Under lower-fertility or moisture-limited growing
conditions, while productivity would be reduced, the relative
performance of the native legumes compared to the cultivated
grain legumes is expected to be improved. These results are
also especially exciting as they are based on only one accession
of each taxa; undoubtedly there is substantial capacity to explore
germplasm for greater productivity, larger seed size and, especially
in wider-spread species, a range in phenological development.
Further collections are needed before variation within species can
be explored fully. The seven most promising species identified
here are also perennials with prospects as forage plants,5,6,10 and
hence could be utilised as dual-purpose crop options where, due
to additional benefits for livestock or farming systems, grain yields
of only 40–60% can compare profitably with annual cultivars.13
It is unlikely that an alternative native Australian legume crop
will replace current grain legumes where they perform well,
but domestication of species with existing adaptation to the
challenging climate and soil conditions may prove a viable option
where current crops are not sustainable.
ACKNOWLEDGEMENTS
This research was funded by Australia’s Rural Industries Research
and Development Corporation (RIRDC). Our thanks to Dr Dai Sutter
and George Weston Technologies, Enfield, New South Wales, for
partnership in the research and useful advice about needs of
the end user. The technical skills of Ms Sabrina Tschirren are
gratefully acknowledged for growing and harvesting the plants in
the glasshouse study.
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