Growth, Yield and Seed Composition of Native Australian Legumes with Potential As Grain Crops

Home , Lupinus angustifolius, Trigonella, Trigonella suavissima

The University of Notre Dame Australia ResearchOnline@ND

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: https://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 ResearchOnline@ND at https://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)andLotus 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 examining approaches to integrate perennial pastures and tree The performance of many exotic legumes used in Australian crops into agricultural systems to reduce dryland salinity. Perennial agriculture is constrained by variable climatic conditions and grain crops have been proposed as an alternative that brings infertile soils.1 Many indigenous Australian legumes may be many sustainability benefits to agriculture while maintaining grain more productive than exotic species when grown under low- production.11,12 Perennial grain crops may only require grain yields fertility or drought conditions.2–5 For example, under low soil P of 40–60% of current crops if they could grow in areas where concentrations the native legumes Kennedia prorepens F. Muell. current crops are less profitable or if they could provide additional 13 and Lotus australis Andrews produced 32 times and 11 times more grazing for livestock. Many of Australia’s native legumes have 6 biomass than lucerne (Medicago sativa), respectively.4 Despite potential as forage species and hence could be used for the dual theirinherentadaptationtochallengingenvironments,thediverse purposes of grain and grazing. legume flora of Australia has received limited assessment for agricultural potential, and focus to date has been on their potential 5–10 as pastures. Few studies investigate Australia’s legumes for ∗ Correspondence to: Lindsay W Bell, CSIRO Ecosystem Sciences, PO Box 102, their potential as alternative grain crops. Furthermore, Australia Toowoomba, QLD 4350, Australia. E-mail: [email protected] has native legumes closely related to globally important grain † legume crops such as Trigonella (fenugreek), Vigna (mungbean), University of Notre Dame Australia, Fremantle, WA 6959, Australia. andGlycine,whichincludesthemostwidelygrownoilseedlegume: a CSIRO Ecosystem Sciences, Toowoomba, QLD 4350, Australia soybean (Glycinemax L.). As well as the potential to provide a wider varietyofadaptedlegumesformodernfarmingsystems,Australian b School of Plant Biology and Institute of Agriculture M081, University of Western legume germplasm in these genera could provide plant breeders Australia, Crawley, WA 6009, Australia across the world with a valuable resource for increased adaptation c CSIRO Ecosystem Sciences, Wembley, WA 6913, Australia to water-limited and infertile environments. Many Australian herbaceous legumes are also perennial and d CentreforLegumesinMediterraneanAgriculture(CLIMA),UniversityofWestern offer sustainability benefits for agriculture. Ongoing work is Australia, Crawley, WA 6009, Australia

J Sci Food Agric (2011) www.soci.org 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 Life cyclea

Native legumes Cullen australasicum (Schltdl.) J. W. Grimes SA44380 Erect sub-shrub A Cullen cinereum (Lindl.) J. W. Grimes AusTrCF 320112 Erect sub-shrub P Cullen graveolens (Domin.) J. W. Grimes AusTrCF 320184 Erect A Cullen tenax (Lindl.) J. W. Grimes AusTrCF 320110 Prostrate to erect P Glycine canescens F. J. Herm NIND001 Perennial twinning P Glycine sp.b NIND004 Perennial twinning ? Glycyrrhiza acanthocarpa (Lindl.) J. M. Black C2N01GA Semi-prostrate to ascending P Kennedia coccinea Vent. NS-26828 Twining P Kennedia prorepens F. Muell. NS-30323 Prostrate to ascending P or A Lotus cruentus Court NF003 Prostrate to ascending P Rhynchosia minima (L.) DC NF013 Prostrate or twining P Swainsona canescens (Benth.) F. Muell. KIMS003 Prostrate or erect, spreading A Swainsona colutoides F. Muell. NIND006 Erect A or P Swainsona kingii F. Muell. NF002 Prostrate or ascending A or P Swainsona purpurea (A. T. Lee) Joy Thomps. KIMS004 Erect to spreading P Swainsona swainsonioides (Benth.) J. M. Black KIMS005 Spreading ascending A Trigonella sauvissima Lindl. Fortescue collection Erect to prostrate or ascending A Commercial grain legumes Cicer arietnum L. (chickpea) Rupali Spreading ascending A Pisum sativum L. (ﬁeld pea) Kaspa Twining, ascending A Lupinus angustifolius L. (narrow-leaf lupin) Mandelup Erect A

a A, annual; P, perennial. b Speciesisunknown.

Although Australian legumes were consumed by Aboriginals, and Trigonella suavissima Lindl. One previous study found that there is no evidence of any kind of their domestication. The use Hardenbergia violacea (Schneev.) Stearn, Crotalaria cunninghamii, of seeds as a food source is mainly documented for several Acacia and Kennedia nigricans Lindl. were worthy of further investigation species,14,15 though there is evidence of Swainsona galegafolia as grain crops, as they possessed relatively large seeds (38 mg, (Andrews) R. Br. (smooth Darling pea) being eaten fresh and 38 mg, and 16 mg, respectively) and contained crude protein tasting similar to common garden pea.16 There are few published content of 210–280 g kg−1 fresh weight.19 However, there is records of the ethnobotany of many Australian native legumes; the little information on the grain or seed production potential ephemeral growth of many species or their rarity in the vegetation and seed composition of many native Australian legumes. may have rendered them an unreliable food source. Antinutritional Therefore, this investigation was conducted to compare the compounds or toxins (e.g. alkaloids, furanocoumarins, and growth characteristics, reproductive development, seed yield, and hydrogen cyanide), present in many Australian legumes, may also seed composition of selected species with those of commercially lower their palatability or edibility. ‘Dilly bags’ (leaching baskets) grown grain legume crops. We found that several native legumes were commonly used by Aborigines to remove toxins by soaking had similar reproductive development, reproductive allocation, certain seeds and legumes in running water for hours or days. and seed composition, but smaller seed size than commercial Despite known toxins in Australian legumes, many secondary grain legume cultivars. 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 METHODS systematically evaluated 14 genera for their adaptation to arid Experimental design and management and winter-dominant semi-arid climatic regions, as well as Seventeen native Australian legumes and three commercial characteristics such as pod indehiscence, growth habit, grain grain legumes (Table 1) were grown in a glasshouse from mid size, seed composition, and likely presence of antinutritional May to December 2008 at the University of Western Australia, ◦ ◦ compounds or toxins.17 This review found that a number Crawley, Australia (31 59 S, 115 53 E). This period was chosen of herbaceous species are likely to be adapted to dry and to coincide with the winter–spring growing season of grain crops infertile environments and possess many desirable attributes in southern Australia, including the three commercial species. for domestication as grain crops. These species include Cullen The 17 native species were prioritised based on information in tenax (Lindl.) J. W. Grimes, Crotalaria cunninghamii R. Br., Glycine the literature,17,20 as well as unpublished evaluations (R Snowball canescens F. J. Herm., Glycyrrhiza acanthocarpa (Lindl.) J. M. and S Hughes, private communication). Seed for several high- Black, Kennedia prorepens, Rhynchosia minima (L.) DC., Swainsona priority species identiﬁed by Bell et al.17 was not available (e.g. canescens (Benth.) F. Muell., Swainsona colutoides F. Muell., Crotolaria species), so either lower-priority or other representative

wileyonlinelibrary.com/jsfa c 2011 Society of Chemical Industry J Sci Food Agric (2011) Growth, yield and seed composition of Australian legumes www.soci.org species were evaluated. Seed was obtained from SARDI Genetic at harvest. Plants were harvested 3 weeks after water was withheld Resource Centre, Urrbrae, SA; Australian Tropical Crop and Forage and separated into pods, seeds, and shoot material. Biomass ◦ Resource Centre, Biloela, QLD; Kimseed International, Osborne samples were dried at 70 C for 72 h and seeds were air dried ◦ Park, WA; Nindethana Seed Service, Albany, WA; and from previous and stored at 25 C prior to compositional analysis. Some species collections made throughout the arid and semi-arid regions of did not flower by the end of the experimental period and hence Western Australia. In addition to the benefits for controlling pests only vegetative shoot biomass was measured. Seeds were ground and diseases, information on self-pollination compatibility of these and then analysed by George Weston Technologies (Enfield, NSW, native species is limited, so plants were grown in the glasshouse in Australia) for fat content using fat Soxhlet extraction, crude protein the absence of pollinators (and without hand pollination) to favour by combustion method, and dietary fibre by phosphate buffer species with higher self-compatible pollination – a desirable trait digestion, following the methods described in the Official Methods in a grain crop. of Analysis of AOAC International.21 Seeds of all taxa were scarified, imbibed on dampened filter paper and sown the following day. Six seeds of each species were Statistical analysis sown per pot, in 10 replicate 300 mm black standard pots with Data for each variable measured were analysed by general analysis drainage holes. Pots were filled with potting mix composed of of variance (ANOVA) in Genstat version 10 (Lawes Agricultural 5 : 2 : 3 (v/v) fine composted pine bark : coco peat : coarse sand, Trust, Rothamsted Experimental Station, Harpenden, UK, 2007). with 1 kg m−3 superphosphate, 2 kg m−3 extra-fine limestone, The least significant difference values at P = 0.05 are shown 0.3 kg m−3 potassium sulfate, 0.2 kg m−3 macro mineral trace at the base of each table. Principle components analysis using elements, 1 kg m−3 ammonium nitrate (Agran 34.0), 2 kg m−3 Genstat was also conducted to investigate correlations amongst dolomite (Ca/Mg), and 0.5 kg m−3 ferrous sulfate heptahydrate. reproductive, growth and seed yield components in 15 of the Pots were placed in a glasshouse at ambient temperature and species; two species which had missing data were not included. 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 RESULTS Plant reproductive development and growth habit 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 All indigenous legumes flowered later than the commercial culti- water. Seedlings were thinned to three plants per pot (i.e. 42 vars of chickpea and lupin (Table 2). The fastest-flowering native plants m−2) at 7 weeks after sowing (i.e. 20 June). This density was species were Trigonella sauvissima, Lotus cruentus, Swainsona chosen to match recommended crop density for the commercial kingii, Glycine sp. and Rhynchosia minima (Fig. 2), all which flow- grain legume species grown under field conditions (i.e. 30–50 ered before the field pea cultivar, taking between 74 and 107 days plants m−2). Because all species grown are indeterminant, water after sowing or 1219–1763 degree days (Table 2). The longest was withheld once the majority of pods had reached maturity: species to flower – Glycyrrhiza acanthocarpa – took 202 days or from 26 weeks after sowing for commercial grain legume species more than 3000 degree days before the appearance of the first (i.e. 10 November) and from 31 weeks after sowing for native flower. A number of native species continued to flower until the legumes species (i.e. 15 December 2008), because of slower onset final harvest (212 days after sowing). Cullen australasicum, Glycine of flowering and maturity. In the glasshouse, temperatures were sp., Kennedia coccinia, L. cruentus, R. minima, Swainsona colutoides, ◦ optimal for plant growth (maxima of 20–30 C and minima of S. kingii, S. purpurea,andT. sauvissima had a distinct flowering ◦ 5–15 C) during the vegetative growth period (mid May to late period (68–101 days) and completed flowering and pod set by ◦ August), and increased by up to 5 C after this time (Fig. 1). 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 Plant growth and seed composition similar to chickpea (457 mm), ranging from 245 (S. kingii)to Over the experimental period, observations of plant growth and 528 mm (Swainsona swainsonioides); the exceptions were the reproductive development were made every 2–3 days, including climbing–twining Glycine and Kennedia, which were staked and days to emergence, days to appearance of first flower, duration hence their height at harvest was greater. The degree of branching of flowering and podding, pod dehiscence (shattering) rating, varied significantly between the species, with T.sauvissima,Glycine number of primary and secondary branches at harvest, and height sp., G. canescens, S. colutoides,andS. swainsonioides exhibiting apical dominance similar to field pea. Only two native species (R.

35 minima and K. prorepens) branched more than chickpea, which is renowned for its highly branched ascending habit. Pod-shattering 30 C) ° scores indicated that four of the native legumes had full seed 25 retention and ﬁve exhibited moderate pod shattering (Table 2). 20

15 Seed yield and yield components

10 Field pea had the highest seed yield and plant biomass of the legumes grown. Glycine sp. yielded more seed and biomass than

Glasshouse temperature ( 5 chickpea, while the seed yield of six other native legumes were 0 50–75% of chickpea and 20–30% of ﬁeld pea (Table 3). Some 11 May 1 Jun 22 Jun 13 Jul 3 Aug 24 Aug 14 Sep 5 Oct 26 Oct of the native species also had a harvest index similar to that of Figure 1. Maximum and minimum glasshouse temperatures during the the commercial cultivars; in particular, L. cruentus had one of the time when the native and commercial grain legumes were grown. lowest total plant biomasses and one of the highest seed yields,

J Sci Food Agric (2011) 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 Thermal time Height at Days to Duration of to first flowera harvestb No. branches Shattering ◦ Species first flower flowering (d) ( Cd) (mm) at harvest scalec

Cicer arietnum 60 44 1003 457 25.21 Lupinus angustifolius 74 – 1219 –– 1 Trigonella sauvissima 74 101 1219 444 7.6 3 Lotus cruentus 81 100 1333 340 19.2 2 Swainsona kingii 92 91 1509 245 20.0 ? Glycine sp. 104 79 1707 702 6.2 2 Rhynchosia minima 107 84 1763 497 28.0 ? Pisum sativum 109 28 1797 1077 6.81 Swainsona colutoides 110 83 1815 438 4.5 ? Swainsona purpurea 111 69 1833 463 13.6 2 Swainsona swainsonioides 111 dnf 1833 528 7.6 ? Cullen cinereum 115 dnf 1909 363 15.8 1 Cullen tenax 116 dnf 1927 491 19.0 1 Cullen graveolens 122 dnf 2038 301 19.0 1 Swainsona canescens 130 dnf 2189 425 15.0 2 Cullen australasicum 132 68 2230 330 19.5 1 Glycine canescens 157 dnf 2741 1366 10.6 2 Kennedia prorepens 167 dnf 2967 782 44.0 ? Glycyrrhiza acanthocarpa 202 dnf na 444 14.0 ?

LSD (P = 0.05) 9 7 167 230 7.6 n/a

a Sum of average daily temperature (i.e. base temperature of 0). b Height of both Glycine species and Kennedia prorepens is of staked plants. c 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 Seed composition (C. australasicum, R. minima, K. prorepens, S. swainsonioides, S. Most native legumes had seed composition within the range purpurea,andS. canescens) produced high amounts of plant of the commercial grain legume cultivars (Table 4). The only biomass, but they had low seed yields and hence very low harvest exceptions were R.minima, which had lower crude protein content indices (<0.10). Notably, we observed these species to have low (<220 g kg−1 DW) and two Cullen species, with fat content over pod set in the glasshouse, which may indicate that they require 110 g kg−1 DW, which was higher than all other legumes tested pollinators to facilitate flower fertilisation. here (12–62 g kg−1 DW) (Table 4). Protein content was highest in A clear distinction between the commercial cultivars and T. sauvissima (373 g kg−1 DW), but a number of native legumes the native legumes was their seed size, with field pea and had crude protein greater than 300 g kg−1. Chickpea and field pea chickpea having seeds weighing 260 mg and 189 mg, respectively protein content was lower than most native legumes. Lupin had (Table 3 and Fig. 2). The native legumes with largest seeds were the highest dietary fibre content (476 g kg−1 DW), but the native G. canescens (17 mg) and R. minima (13.3 mg). Seeds of Swainsona Glycine and Lotus species also had high dietary fibre (>300 g kg−1 species were between 2.7 and 5.5 mg, Cullen were around 5 mg DW). The remainder of the native species had fibre content in the and the two smallest seeded species were T. suavissima and L. range 196–286 g kg−1 DW (Table 4). 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 DISCUSSION tested (Fig. 3). We expected that the seed yield, reproductive allocation and seed Principal component analysis showed close correlations be- composition of grain legume cultivars would be far superior to the tween seed mass and seed yield, mainly associated with the native Australian legumes, which have received no domestication commercial cultivars (Fig. 2). Of the major factors represented in or selection for grain yield or quality. However, yields greater than the two first principal components, time to first flower was in- 40% of chickpea and similar reproductive allocation, comparable versely correlated with harvest index and seed number per plant, phenology, and moderate to low pod shattering were found and biomass and plant height were inversely correlated with seeds in Glycine sp., L. cruentus, S. kingii and S. colutoides.Three per plant. Amongst the native legumes, the two Glycine species other species – C. cinereum, C. tenax,andG. canescens – flowered were differentiated based on both their higher seed yield and seed later than the grain legume cultivars (115, 116 and 157 days mass. In addition to the Glycine species, the six native species in after sowing, respectively) but also yielded >40% of chickpea. the top right-hand quadrant were those with the highest seed Universally, seed size was ∼14 times smaller in the native legumes yields and harvest index and more rapid onset of flowering (Fig. 2). than the domesticated species. The highest yields were in species

wileyonlinelibrary.com/jsfa c 2011 Society of Chemical Industry J Sci Food Agric (2011) Growth, yield and seed composition of Australian legumes www.soci.org

4 PC2

1 Pod # C. graveolens C. cinereum Trigonella sauvissima # branches Seed # S. purpurea Lotus cruentus Kennedia prorepens S. canescens C. tenax S. kingii C. australasicum . S. colutoides TT to flower/DTF S. swainsonioides R. minima Seeds per pod PC1 0 -3-2-10123

Glycine canescens Glycine sp. HI -1 Biomass yield Chickpea Height

Seed yield -2 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).

4.0 some potential as perennial grain crops, which could provide y = -0.7538x + 3.1403 a number of environmental and production benefits in farming 2 -1 L. cruentus R = 0.8248 11,12 T. suavissima systems. Overall, we believe that the seven native Australian 3.0 S. kingii C. tenax S. colutoides Glycine sp. legumes mentioned above have adequate growth, yield, and C. cinereum G. canescens seed composition to be subject to further investigation for their 2.0 C. graveolens C. australasium agricultural potential as grain crops. R. minima Field pea The experiment reported here was conducted under glasshouse Chickpea seed number plant conditions with warmer temperatures, and more favourable 10 1.0 moisture and fertility than would be expected in the field. Yet Log the similar plant density, crop productivity, and seed mass in field- 0.0 grown commercial grain legumes provides some basis for these 0.0 1.0 2.0 3.0 results to be translated to field conditions. First, plant density Log seed mass (mg) 10 in pots (i.e. 42 plants m−2) was in the range recommended for Figure 3. Relationship between individual seed mass and seed number commercial grain legumes in the field,22 suggesting that inter- producedperplantfornativeAustralianlegumesandtwocommercialgrain plant competition for light and soil resources was similar to field legume cultivars. Those native species with poor seed set, presumably due conditions. Secondly, harvest index, seed mass, and crop grain and to pollination problems, are omitted. biomass yield calculated on an area basis are also similar to those reported for crops grown under favourable field conditions.22 that produced a large number of seeds per plant; the one Compared to field-grown crops, chickpea in this study had similar − − exception was the largest-seeded native legume: G. canescens. grain yield (1.9 t ha 1 comparedto1.5–2.0tha 1 in the field), − − Many of the seven native legumes with the highest reproductive crop biomass (3.3 t ha 1 compared to 3–5.7 t ha 1 in the field), allocation and grain yield in this study are also perennials (i.e. and seed mass (189 mg per seed compared to 156–186 mg per Glycine sp., L. cruentus, C. tenax and G. canescens) and hence offer seed in the field), though harvest index was higher (0.6 compared

J Sci Food Agric (2011) c 2011 Society of Chemical Industry wileyonlinelibrary.com/jsfa www.soci.org 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 Seed yield Total biomass HI Seed mass Seed no. Pods per Seeds Species (g per plant) (g per plant) (g g−1) (mg per seed) per plant plant per pod

Pisum sativa 9.94 20.40.50 258.939123.2 Glycine sp. 4.78 8.9 0.54 11.2 428 79 5.4 Cicer arietnum 4.56 7.70.60 188.724280.9 Lotus cruentus 3.40 5.3 0.65 1.5 2320 184 12.6 Cullen tenax 2.77 8.7 0.30 5.2 515 515 1.0 Glycine canescens 2.68 7.8 0.35 16.9 159 38 4.2 Swainsona kingii 2.18 4.6 0.47 2.7 809 103 7.9 Cullen cinereum 2.06 6.6 0.30 5.2 393 393 1.0 Swainsona colutoides 2.01 9.6 0.21 3.1 645 66 9.8 Trigonella sauvissima 1.43 5.0 0.35 1.2 1185 340 3.5 Cullen australasicum 1.42 15.6 0.09 8.7 177 177 1.0 Rhynchosia minima 0.86 13.4 0.07 13.3 67 43 1.5 Cullen graveolens 0.82 6.1 0.11 5.7 141 141 1.0 Swainsona canescens 0.17 10.0 0.02 2.7 53 17 3.1 Swainsona purpurea 0.04 10.6 0.004 3.2 14 4 3.5 Kennedia prorepens 0.02 13.7 0.001 6.3 1 0.6 1.6 Swainsona swainsonioides 0.01 13.6 0.001 5.5 3 4 0.7 Glycyrrhiza acanthocarpa n.a 5.2 n.a n.a – – – LSD (P = 0.05) 0.94 2.5 0.09 17.7 210 120 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 Crude protein (g kg−1 DW) Fat (g kg−1 DW) Dietary Fibre (g kg−1 DW)

Trigonella sauvissima 373 53 204 Lupinus angustifolius (lupin) 369 51 476 Cullen cinereum 362 118 262 Swainsona kingii 343 25 236 Glycine canescens 340 62 317 Swainsona swainsonioides 325 36 196 Cullen australasicum 324 38 286 Glycine sp. 322 52 350 Cullen tenax 321 113 279 Lotus cruentus 320 59 429 Cullen graveolens 304 57 n.a. Swainsona colutoides 275 21 239 Swainsona canescens 269 40 227 Pisum sativa (ﬁeld pea) 263 12 154 Glycyrrhiza acanthocarpa 261 n.a. n.a. Cicer arietnum (chickpea) 229 45 257 Rhynchosia minima 210 n.a. n.a.

n.a., insufﬁcient seed for analysis.

to 0.4–0.5 in the field). Field pea grown in our experiment seemed These comparisons suggest that under favourable field growing to convert biomass to grain yield more efficiently here than under conditions the best native legumes included in our glasshouse field conditions; grain yield was 4.2 t ha−1 compared to 1.5–2.7 t experiment could yield 1.2–2.0 t ha−1 of grain and 2.4–4.0 t ha−1 ha−1 in the field and harvest index was 0.5 compared to 0.35–0.4 in of biomass in the field, with similar seed size but possibly reduced the field, but seed mass and total crop biomass were similar to field harvest index. Under more arid or infertile growing conditions observations (259 mg per seed compared to 179–240 mg per seed these productivity levels would be reduced, but the expected and 8.5 t DM ha−1 here compared to 6–10 t ha−1, respectively).22 relative reduction in productivity of native legumes would be

wileyonlinelibrary.com/jsfa c 2011 Society of Chemical Industry J Sci Food Agric (2011) Growth, yield and seed composition of Australian legumes www.soci.org less than for the exotic legumes.3–5 The low seed set in some in this accession of C. cinereum.7 The T. suavissima accession tested species (S. canescens, S. purpurea, S. swainsonoides, K. prorepens, here flowered more quickly (74 days) than reported previously C. australasicum,andR. minima) is likely to be due to the lack for this species (111–118 days),17 suggesting that material that of pollinators in the glasshouse and confirms the dominance flowers much earlier may be available. In this study, R. minima of open pollination suspected in these species.17 Similarly, the flowered after 107 days, which is midway between that reported photoperiod sensitivity of many native legumes is unknown and previously (43–142 days).26 Overall, there have been few studies the photoperiod regime during the experimental period may have of variation in agronomic traits of native Australian legumes, but influenced the growth and development of some species. For current evidence suggests there is substantial variation that could example, long days (>12–13 h) are thought to induce flowering be exploited to improve productivity, seed size, and reproductive in C. australasicum,7 which may explain the long period to the start development to fit into different environments.1 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 CONCLUSION this study suggest that higher yields still might be expected when This preliminary investigation of a range of native Australian sown in spring. legumes, chosen for this study because of adaptation to An important consideration for the potential domestication arid and semi-arid winter dominant rainfall conditions, has of native legumes is the end use of the grain, in particular, shown that, while many species had much lower yields and concentration of protein, fats/oils, fibre and the presence of reproductive allocation, seven species warrant further evaluation antinutritional or beneficial compounds. In this study we found for domestication for agriculture. These seven species exhibited that most of the native legumes had higher grain crude protein grain yields of 40–60% of cultivated grain legumes with grain content than field pea and chickpea, and some were similar to the protein, fat and fibre in the range desirable in food and highproteincontentfoundinlupin.Previousstudieshavereported feed industries. Under lower-fertility or moisture-limited growing 19,23 lower grain protein contents in native Australian legumes, conditions, while productivity would be reduced, the relative though it is unclear whether this was calculated on a dry mass performance of the native legumes compared to the cultivated basis. Grain of the two Cullen species had a higher fat content grain legumes is expected to be improved. These results are −1 (>110 g kg DW) than the other native legume seeds tested also especially exciting as they are based on only one accession −1 here, and fat contents greater than 80 g kg werefoundintwo of each taxa; undoubtedly there is substantial capacity to explore 19 species not tested here: K. nigricans and H. violacea. Cullen seed germplasm for greater productivity, larger seed size and, especially has an adherent pod so fats and oils could be concentrated there, in wider-spread species, a range in phenological development. although, surprisingly, the inclusion of this pod in the analysed Further collections are needed before variation within species can sample did not result in elevated fibre content. Despite the high be explored fully. The seven most promising species identified oil content in Cullen, these levels are much lower than found here are also perennials with prospects as forage plants,5,6,10 and ∼ −1 in soybean ( 200 g kg ) and other oilseed crops. A number hence could be utilised as dual-purpose crop options where, due of Australian legumes are also known to produce toxins (e.g. to additional benefits for livestock or farming systems, grain yields swainsonine in some Swainsona species, hydrogen cyanide in of only 40–60% can compare profitably with annual cultivars.13 Australian Lotus) and other bioactive compounds (e.g. isoflavones, It is unlikely that an alternative native Australian legume crop especially phytoestrogens in Glycine, furanocoumarins in Cullen) will replace current grain legumes where they perform well, which could limit their use in food products. However, a number but domestication of species with existing adaptation to the of these compounds are pharmaceutically useful and could offer challenging climate and soil conditions may prove a viable option 17 potential as natural medicines. Many cultivated grain legumes where current crops are not sustainable. 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 ACKNOWLEDGEMENTS within the taxa is required. This research was funded by Australia’s Rural Industries Research In this study we only compared one accession of each species, and Development Corporation (RIRDC). Our thanks to Dr Dai Sutter yet there could be substantial genetic variation in key agronomic and George Weston Technologies, Enfield, New South Wales, for traits within species, and much more productive genotypes are partnership in the research and useful advice about needs of likely to exist. This is clearly shown by Snowball et al.,20 who found the end user. The technical skills of Ms Sabrina Tschirren are that seed yield and plant survival in the field varied considerably gratefully acknowledged for growing and harvesting the plants in among accessions of native species, including K. prorepens the glasshouse study. 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 REFERENCES (16.9 mg) than recorded previously for this species (5.9–8.9 mg).17 1 Thomson BD, Siddique KHM, Barr MD and Wilson JM, Grain legume species in low rainfall Mediterranean-type environments I. Others have shown that seed size varies by two–four times within Phenology and seed yield. Field Crops Res 54:173–187 (1997). 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