Intra-fruit seed abortion in a wind dispersed tree,Dalbergia sissoo Roxb: Proximate mechanisms

Dalbergia sissoo, a tropical tree with wind-dispersed pods, exhibits a highly positively skewed distribution of seeds per pod with predominantly only one of the four or five ovules maturing into seed. The abortion cannot be attributed to lack of pollen or resources. This study examines the hypothesis that the abortion is due to an intense rivalry among the developing sibs to gain dispersal advantage. Aqueous extract of the dominant embryos at the stigmatic end that generally develop to maturity significantly inhibited the uptake of labelled sucrose by the young developing (subject) embryos in an in vitro assay system. Extracts of tissues such as subordinate (peduncular embryos), unfertilized ovules and pod coat did not cause such inhibition. Aqueous diffusate of dominant embryos also inhibited the uptake of labelled sucrose by subject embryos. The chemical substance responsible for the inhibition appears to be heat-stable and non-proteinaceous. HPLC analysis indicated the presence of two retention time peaks, different from that of standard indole acetic acid, but with considerable overlap. We hypothesize that the compound could be an indole derivative. We propose that the stigmatic embryos have a head start due to earlier fertilization and produce a chemical that either directly (by metabolically killing) or indirectly (by preventing the uptake of assimilates) kills the proximally placed peduncular embryos.     

Single-Seeded Fruits and Seedling Establishment in Dalbergia miscolobium Benth. (Papilionaceae)1

Dalbergia miscolobium (Papilionaceae) is a tree of the Brazilian cerrado that produces ovaries with two ovules. Only one ovule usually matures, but when two mature, they are smaller than seeds of single-seeded pods. Germination percentage was lower in small seeds than in medium or large seeds. In general, small seeds gave rise to smaller seedlings (dry mass) than did medium and large seeds. The root/shoot ratio, however, was very similar for the first six months, indicating that this ratio is consistent across seedling sizes. There was no correlation between seed size and relative growth rate. Single-seeded fruit production in D. miscolobium resulted in greater seed reserves and increased fruit dispersal distance; single-seeded fruits were lighter than double-seeded ones. Together, these two features should increase the probability of successful seedling establishment of seeds from single-seeded versus double-seeded fruits.     

Selective embryo abortion in a perennial tree-legume: a case for maternal advantage of reduced seed number per fruit

In this study, I analyzed time-course of embryo abortion, positional bias in seed maturation and maternal costs of seed packaging in Cercis canadensis. While basal embryos experience similar rates of abortion as those in other positions during the first week of development, abortion rates peak during the second week. Head start in resource sequestration by stigmatic embryos may explain high rates of basal embryo abortion. Similar seed packaging costs and seed mass for single and multi-seeded pods suggests that maternal parent regulates pod size in accordance with seed numbers per pod investing equally in the surviving offspring. Competition during early developmental period results in the abortion of less competitive embryos allowing for optimal resource investment.     

Seed abortion in wind-dispersed pods of Dalbergia sissoo: maternal regulation or sibling rivalry?

Summary Dalbergia sissoo, a wind-dispersed tropical tree, shows a positively skewed distribution of seeds per pod. This is attributed to the enhanced dispersal advantage of few-seeded pods due to their reduced wing loading (ratio of weight to pod surface area) and low settling velocity. The proximate mechanisms causing the positively skewed distribution were investigated. The distribution could not be attributed to the distribution pattern of ovule number per ovary, pollen grain limitation, lack of ovule fertilization, or post-fertilization elimination of many-seeded pods. Rather, it was caused by the post-fertilization abortion of seeds within a pod 2 weeks after fertilization. This intra-pod seed abortion (IPSA) is due to a dominance hierarchy of fertilized ovules from the distal (near stigma) to the basal end, generated by the temporal differences in fertilization. The dominant developing seeds at the distal end cause the abortion of others through the production and diffusion of an aborting agent. When the dominance hierarchy of the siblings is not intense, pods are formed with more than one seed. We argue that the positively skewed distribution of seeds per pod is not due to maternal regulation but is a result of sibling rivalry. We propose that this sibling rivalry is generated by genetic differences in pollen grain fitness and disucss the results in the context of parent-offspring conflict.     

Intensity of pre‐dispersal seed predation in the invasive legume Leucaena leucocephala is limited by the duration of pod retention

Abstract The intensity of seed predation the invasive tropical legume Leucaena leucocephala by the bruchid Acanthoscelides macropthalmus was investigated in south‐eastern Queensland, Australia. The number of seeds damaged by A. macropthalmus as a proportion of total seeds available was found to increase the longer the pods remained on the tree. Seed predation ranged from a mean of 10.75% of seeds on pods that remained on the plant for 1 month and increased to 53.54% for pods that remained of the plant for 4 months. The low bruchid populations at high pod densities results in ‘predator satiation’. However, pods dehisce over time and the proportion of pods available over time to the bruchid correspondingly declines. By the time bruchid densities build up, most pods have dehisced and the seeds consequently escape predation. As a result the number of seeds lost to bruchid damage increases only marginally over time. Despite the levels of seed predation observed over the course of the study, the number of seeds in the soil seedbank almost doubled over time increasing from 8.5 seeds m^?3 to 15.5 seeds m^?3 over a 4‐month period. Levels of seed predation and addition of seeds to the soil seedbank were not correlated. The taxonomic (subspecies) status and apparency of host plants as measured by plant and patch traits (average plant height, density of podding plants and patch size) did not influence levels of seed predation. Pre‐dispersal seed predation studies need to take into account the pod/seed retention behaviour of the plant. The ability of the bruchid to regulate the invasiveness of Leucaena through influencing its demography is likely to be diminished if the insect populations cannot increase rapidly enough to use the seeds before pod dehiscence.     

Patterns of fruit and seed production inBauhinia ungulata (Leguminosae)

Patterns of seed and fruit production ofBauhinia ungulata, a small tree legume indigenous in tropical America, were studied in Costa Rica. Only about 8% of flowers produced fruits. The average pod had 19 ovules and about two thirds of these began seed development, with mature pods containing an average of 9.7 mature undamaged seeds. About half of the mature pods were damaged by herbivores and within these, 27% of ovules or seeds had been eaten. Among trees there was no significant variation in pod production, but the number of ovules per pod and seed production per pod varied significantly. Within infructescences most pods were retained at middle positions. Within pods, the probability of an ovule developing into a seed increased toward the distal end. The pattern of seed and fruit production in this species agrees well in general with that reported for other neotropical legumes. The abortion of seeds and fruits can be regarded as a way of controlling maternal investment, and as a response to herbivory.     

Selenium Hyperaccumulator Plants Stanleya pinnata and Astragalus bisulcatus Are Colonized by Se-Resistant, Se-Excluding Wasp and Beetle Seed Herbivores

Selenium (Se) hyperaccumulator plants can concentrate the toxic element Se up to 1% of shoot (DW) which is known to protect hyperaccumulator plants from generalist herbivores. There is evidence for Se-resistant insect herbivores capable of feeding upon hyperaccumulators. In this study, resistance to Se was investigated in seed chalcids and seed beetles found consuming seeds inside pods of Se-hyperaccumulator species Astragalus bisulcatus and Stanleya pinnata. Selenium accumulation, localization and speciation were determined in seeds collected from hyperaccumulators in a seleniferous habitat and in seed herbivores. Astragalus bisulcatus seeds were consumed by seed beetle larvae (Acanthoscelides fraterculus Horn, Coleoptera: Bruchidae) and seed chalcid larvae (Bruchophagus mexicanus, Hymenoptera: Eurytomidae). Stanleya pinnata seeds were consumed by an unidentified seed chalcid larva. Micro X-ray absorption near-edge structure (µXANES) and micro-X-Ray Fluorescence mapping (µXRF) demonstrated Se was mostly organic C-Se-C forms in seeds of both hyperaccumulators, and S. pinnata seeds contained ∼24% elemental Se. Liquid chromatography–mass spectrometry of Se-compounds in S. pinnata seeds detected the C-Se-C compound seleno-cystathionine while previous studies of A. bisulcatus seeds detected the C-Se-C compounds methyl-selenocysteine and γ-glutamyl-methyl-selenocysteine. Micro-XRF and µXANES revealed Se ingested from hyperaccumulator seeds redistributed throughout seed herbivore tissues, and portions of seed C-Se-C were biotransformed into selenocysteine, selenocystine, selenodiglutathione, selenate and selenite. Astragalus bisulcatus seeds contained on average 5,750 µg Se g⁻¹, however adult beetles and adult chalcid wasps emerging from A. bisulcatus seed pods contained 4–6 µg Se g⁻¹. Stanleya pinnata seeds contained 1,329 µg Se g⁻¹ on average; however chalcid wasp larvae and adults emerging from S. pinnata seed pods contained 9 and 47 µg Se g⁻¹. The results suggest Se resistant seed herbivores exclude Se, greatly reducing tissue accumulation; this explains their ability to consume high-Se seeds without suffering toxicity, allowing them to occupy the unique niche offered by Se hyperaccumulator plants.     

Seed development in Pisum

The Ih and lh _i alleles have been shown previously to reduce the level of endogenous gibberellin A₁ (GA₁) in shoots of pea (Pisum sativum L.), resulting in a dwarf phenotype compared with the wild type, cv. Torsdag (Lh). In addition, plants homozygous for the lh ⁱ allele have reduced seed yield compared with Lh (tall, wild type) and lh (dwarf) plants. In this paper we show that the lh ⁱ mutation is expressed in developing seeds and pods. Comparison of GA levels in young shoots and developing seeds of genotypes lh and lh ⁱ demonstrates that the relative severity of the two mutations varies in different tissues. Homozygous h ⁱ seeds have reduced GA levels, weigh less, and are less likely to develop to maturity when compared with Lh seeds. However, fertilization of lh ⁱ plants with Lh pollen increases seed GA levels, seed weight and seed survival, indicating that an increase in seed GA levels due to the presence of the Lh allele can restore normal seed growth. Pods developing on self-pollinated lh ⁱ plants are shorter than pods on Lh (wild type) plants, although this may be an indirect effect of the increased seed abortion of lh _i plants. Based on these results we suggest that endogenous GAs play an important role in the development of seeds of P. sativum L.Abbreviations GA_(n) gibberellin A_n We wish to thank Katherine McPherson, Peter Newman, Leigh Johnson and Peter Bobbi for technical assistance, Professor L. Mander (ANU, Canberra) and Professor B.O. Phinney (UCLA, USA) for labelled GA standards, and the Australian Research Council for financial support.     

Stigmatic inhibition of pollen grain germination-its implication for frequency distribution of seed number in pods of Leucaena leucocephala (Lam) de Wit

Summary Seed number per pod is negatively skewed in Leucaena leucocephala. Post-fertilisation abortion of pods does not completely explain the skewed distribution. It is shown that a pre-fertilisation regulation effectively restricts the formation of pods with less than a certain number of seeds. The regulation is rendered through the inhibition of pollen grain germination by the stigmatic fluid when the pollen grain is less than a certain critical load in the stigma. Such a control is shown to have implications in the evolution of pollen package units, generation of male gametophytic competition and the economy of packing seeds in pods.     

种子植物的选择性败育及其进化生态意义

种子植物的选择性败育是指植株在花粉源、传粉次序、果实在植株上的位置和发育果实中的种子数目等因素或者这些因素综合作用的基础上对发育中的幼果或种子选择性败育的现象。植株可以选择性地败育位于果序顶部或基部的果实以及位于果实基部、中部或柱头端的种子。此现象在被子植物中比较普遍,特别是在豆科、十字花科和紫草科中最为常见。导致植物选择性败育的主要原因主要有资源限制和遗传因子两个方面。植物通过选择性败育部分自交或基因型较差的果实或种子,不仅可以提高母本和后代的适合度,而且还可以提高果实或种子的扩散效率。因此,对选择性败育的研究在深入了解植物的结实结籽格局、探讨其进化式样与机制等方面具有重要意义。该文系统总结了国际上有关植物选择性败育的研究工作,重点介绍了选择性败育发生的式样、导致选择性败育的因素、选择性败育的进化生态意义,以及目前研究选择性败育现象的主要方法,并对该领域今后研究前景进行了展望。     

Application of a probabilistic model for analysing the abortion of seeds and pods in winter oilseed rape (Brassica napus)

In winter oilseed rape (WOSR), only a subset of ovules can develop into seeds in the majority of pods. Any difficulty during the process of seed production may result in seed or pod abortion. This study aimed to reproduce the process of seed development in WOSR based on a limited number of parameters. As a result of the complexity of the developmental patterns of WOSR, it is challenging to identify the roles of various factors that influence seed production using an experimental approach. Here, we present a stochastic probabilistic model of seed development. The generalised least squares method was implemented to estimate the model parameters using the experimental data. Experiments were done in Grignon (France) in 2008 and 2009. The variations in the parameters were analysed according to the following four factors: year, pod rank, inflorescence position and ramification‐clipping treatment. The year had no effect on the number of ovules per ovary (μ) and the probability of seed viability (p). The proportion of effective pollen grains (k) significantly decreased with pod rank at the end of the main stem. Inflorescence position influenced the number of ovules per ovary (μ: 30.8–33.8 from top to bottom) and the parameter k. The mean number of seeds per pod on the main stem and the bottom ramification along the stem was larger than the other ramifications within one plant. Ramification‐clipping treatment increased the number of ovules per ovary (μ: 31 for control plants and 32 for clipped plants), the parameter k and the number of seeds per pod (p). This effect could be due to the competition for assimilates between the pods and seeds. Furthermore, the distribution parameters of the pollen number per stigma (m) remained stable, and the probability of pod survival (Bo) varied with different factors, including the year, pod rank and inflorescence position. Our results indicate that pollen germination is a factor that determines final seed number. This model can identify the impact of each of the factors that lead to the abortion of seeds and pods in WOSR, which include the position effect, assimilate competition and pollination limitation. However, further studies on the pollination process in WOSR should help to refine this model.     

EFFECT OF OVULE POSITION IN THE POD ON PATTERNS OF SEED FORMATION IN TWO SPECIES OF LATHYRUS (LEGUMINOSAE: PAPILIONOIDEAE)

Using a combination of observations of fate of ovules in matured fruits and of fluorescence techniques to study pollen tube growth and fertilization of ovules, we examined patterns of seed formation within pods in natural populations of two species of Lathyrus, L. sylvestris and L. latifolius. We also examined variation in these patterns within and among populations and between two consecutive years. In both species, only a portion of the ovules were fertilized. Fertilization occurs over a period of several days and ovules at the stigmatic end of the fruit are the first to be fertilized. Fertilized ovules farthest from the stigma are closest to the maternal nutrition. The pattern of embryo abortion is interpreted as a balance between the early start and genetic quality of embryos near the stigma, on the one hand, and the nutritional advantage of proximity to maternal nutrients, on the other. Differences between the two species in patterns of seed maturation are postulated to be related to differences in breeding system. In L. sylvestris, a higher frequency of selling leads to less genetic diversity of pollen deposited on the stigma, lower competition among potential sires, and a more nearly random pattern of ovule fertilization and maturation within the pod.     

Partitioning of 14C-photoassimllates within seeds of Phaseolus coccineus (Lam.) and Phaseolus coccineus x Phaseolus vulgaris L.

The objective of this study was to determine partitioning within seeds of ¹⁴C-photoassimilates at three stages of seed development in two Phaseolus crosses — P. coccineus Lam. selfed, and P. coccineus x P. vulgaris L. Abortion of the interspecific embryos occurred when the seed reached 10 mm seed length. When expressed as sink strength (% dpm) or sink activity (% dmp/d.wt.) there were no differences in partitioning of ¹⁴C-photoassimilates when whole seeds were analyzed. If the seed was divided into seed coat, liquid endosperm, and embryo, the sink activity of the interspecific embryo was higher than that of the embryo in the selfed seed. Therefore, abortion of these interspecific Phaselus embryos appeared not to be caused by a lack of photoassimilates.Assistant Professor, Professors, respectively.Contribution from the Agr. Expt. Station, University of Minnesota, St. Paul, MN 55108. Paper No. 13,548, Scientific Journal Series. This research was supported in part by the Science and Education Administration of the United States Department of Agriculture under Grant 59-2271-9-2-020-0 and in part by a grant from the Minnesota Soybean Research and Promotion Council.     

Impairment in reproductive development is a major factor limiting yield of black gram under zinc deficiency

Black gram [Vigna mungo (L.) Hepper] cv. IPU 94 plants grown in sand culture with deficient zinc (0.1 μM Zn) nutrition and those deprived of normal (1 μM) Zn supply at the initiation of flowering, showed decrease in dry matter production and especially seed yield. These plants showed a decrease in the size of anthers and stigmatic heads, pollen producing capacity of the anthers and stigmatic exudations. Zn deficiency caused structural alterations in exine and retarded germination of pollen grains and tube growth. The pollen extracts and stigmatic exudates of the Zn-deficient plants showed increase in activity of acid phosphatase isoforms and inhibition of esterase isoforms. Zn deficiency led to decrease in number of pods, seeds per pod and seed mass, altered seed coat topography and reduced seeds germinability. Low seed yield under Zn deficiency is attributed to a role of Zn in pollen function, as also in pollen-pistil interaction conducive to fertilization and development of seeds.     

Changes in Baboon Feeding Behavior: Maturity-dependent Fruit and Seed Size Selection within a Food Plant Species

Despite considerable inter- and intraindividual variation in fruit and seed size in many plant species, researchers have given little attention to the relevance of the traits for primate fruit choice within a food plant species and its implications for tree regeneration. We studied feeding behavior and selectivity of olive baboons (Papio anubis) in the African locust bean (Parkia biglobosa, Mimosaceae), via direct observations of habituated groups and indirect evidence from leftovers of pods after feeding events. Olive baboons acted as both seed predators and dispersers for Parkia biglobosa. They fed on and destroyed unripe seeds, and swallowed intact ripe seeds when consuming mature fruit pulp. Predation rate was high, and only 10% of the seeds were dispersed. Predation and dispersal of seeds is linked to seed number and size. Digestible unripe seeds accounted for 10% of the unripe fruit mass, while indigestible ripe seeds made up 28% of the mature fruit mass. With these constraints, olive baboons increased food gain per fruit by selecting unripe pods containing a high number of large and heavy seeds. Consequently, only pods with fewer and smaller seeds remained for maturation. Thereafter, baboons fed on mature pods containing the smallest seeds, and exploited pods with more seeds to a greater extent than those with fewer seeds. Thus, fruits with small seeds and an intermediate seed number contributed the most to dispersal by baboons.     

森林鼠类对秦岭南坡3种壳斗科植物种子扩散的影响

种子作为森林鼠类的主要食物来源,它们的扩散和更新很大程度上依赖于鼠类的传播。在鼠类扩散种子的过程中,种子特征和食物相对丰富度是影响鼠类对种子进行何种选择策略的重要因素。2011—2012年的8—12月,采用塑料片标记法在秦岭南坡的佛坪国家级自然保护区内调查了森林鼠类对同域分布的3种壳斗科植物(锐齿槲栎Quercus aliena var.acuteserrata、栓皮栎Q.variabilis和短柄枹栎Q.serrata var.brevipetiolata)种子的扩散差异。结果表明:(1)鼠类倾向于贮藏营养价值较大的栓皮栎种子,并且其贮藏距离也最远(2011:1.52 m,2012:4.03 m),3种种子在食物相对丰富度较低年份被贮藏的距离均较远。(2)在食物相对丰富度较高的年份(2011),种子的消耗速率较慢,在种子释放10 d后种子释放点仍有67.33%的种子,贮藏量较高,至实验结束仍有29.67%的种子被贮藏。在食物相对丰富度较低的年份(2012),种子消耗速率较快,在种子释放后10 d内所有种子均被取食或搬离种子释放点,贮藏量较低,至实验结束仅有3.83%的种子仍被贮藏。(3)虽然栓皮栎种子的贮藏量最大,被贮藏后的存留量也最大,但其在实验地的分布却较小,说明种子扩散仅是植物分布与存活的第一步。以上结果表明,鼠类倾向于贮藏营养价值高的种子。在食物相对丰富度较高的年份会更多的贮藏种子,但种子被贮藏的距离较近,在食物相对丰富度较低的年份会更多的取食种子。     

Salt sensitivity in chickpea (Cicer arietinum L.): ions in reproductive tissues and yield components in contrasting genotypes

The reproductive phase in chickpea (Cicer arietinum L.) is affected by salinity, but little is known about the underlying cause. We investigated whether high concentrations of Na⁺ and Cl^– in the reproductive structures influence reproductive processes. Chickpea genotypes contrasting in tolerance were subjected to 0, 35 or 50 mm NaCl applied to soil in pots. Flower production and abortion, pod number, percentage of empty pods, seed number and size were evaluated. The concentrations of Na⁺, K⁺ and Cl^– were measured in various plant tissues and, using X‐ray microanalysis, in specific cells of developing reproductive structures. Genotypic variation in reproductive success measured as seed yield in saline conditions was associated with better maintenance of flower production and higher numbers of filled pods (and thus seed number), whereas seed size decreased in all genotypes. Despite the variation in reproductive success, the accumulation of Na⁺ and Cl^– in the early reproductive tissues of developing pods did not differ between a tolerant (Genesis836) and a sensitive (Rupali) genotype. Similarly, salinity tolerance was not associated with the accumulation of salt ions in leaves at the time of reproduction or in seeds at maturity.     

Genetic analyses of agronomic and seed quality traits of synthetic oilseedBrassica napus produced from interspecific hybridization ofB. campestris andB. oleracea

The heritability, the number of segregating genes and the type of gene interaction of nine agronomic traits were analysed based on F₂ populations of synthetic oilseedBrassica napus produced from interspecific hybridization ofB. campestris andB. oleracea through ovary culture. The nine traits—plant height, stem width, number of branches, length of main raceme, number of pods per plant, number of seeds per pod, length of pod, seed weight per plant and 1000-seed weight—had heritabilities of 0.927, 0.215, 0.172, 0.381, 0.360, 0.972, 0.952, 0.516 and 0.987 respectively, while the mean numbers of controlling genes for these characters were 7.4, 10.4, 9.9, 12.9, 11.5, 21.7, 20.5, 19.8 and 6.4 respectively. According to estimated coefficients of skewness and kurtosis of the traits tested, no significant gene interaction was found for plant height, stem width, number of branches, length of main raceme, number of seeds per pod and 1000-seed weight. Seed yield per plant is an important target for oilseed production. In partial correlation analysis, number of pods per plant, number of seeds per pod and 1000-seed weight were positively correlated with seed yield per plant. On the other hand, length of pod was negatively correlated (r = -0.69) with seed yield per plant. Other agronomic characters had no significant correlation to seed yield per plant. In this experiment, the linear regressions of seed yield per plant and other agronomic traits were also analysed. The linear regression equation wasy = 0.074x₈ + 1.819x₉ + 6.72x₁₂ -60.78 (R ² = 0.993), wherex ₈, x₉ and x₁₂ represent number of pods per plant, number of seeds per pod and 1000-seed weight respectively. The experiment also showed that erucic acid and oil contents of seeds from F₂ plants were lower than those of their maternal parents. However, glucosinolate content was higher than that of the maternal plants. As for protein content, similar results were found in the F₂ plants and their maternal parents. It was shown that the four quality traits, i.e. erucic acid, glucosinolate, oil content, and protein content, had heritability values of 0.614, 0.405, 0.153 and 0.680 respectively.     

The Effects of Primates and Squirrels on Seed Survival of a Canopy Tree,Afzelia quanzensis,in Arabuko‐Sokoke Forest,Kenya1

I examined the fate of seeds from ten focal trees of Afzelia quanzensis (Leguminosae), a canopy tree in the Arabuko‐Sokoke, Kenya. The study was conducted for one fruiting season, between August 1990 and July 1991. Yellow baboons (Papio cynocephalus), Syke's monkeys (Cercopithecus albogularis), sun squirrels (Heliosciurus rufobrachium), and bush squirrels (Paraxerus palliatus) were all observed to interact with A. quanzensis seeds at various stages of pod development. Baboons and squirrels consumed high percentages of seeds when they were still immature, but the seeds were still unavailable to Syke's monkeys at this stage. Baboons bit open the hard green pods and squirrels gnawed through the pods to extract the immature seeds (hereafter referred to as seed predation), but monkeys were unable to open the pods. Upon maturity, the pods opened slightly, revealing red arils that were sought by baboons, monkeys, and squirrels. Monkeys removed the highest percentage of mature seeds from these pods. These mammal dispersers ate the arils from the mature seeds and discarded the viable part that germinates (hereafter referred to as seed dispersal). My data indicate that baboons and squirrels are seed predators while monkeys are seed dispersers of A. quanzensis.     

Genetic load,nutrient limitation,and seed production in Lupinus texensis (Fabaceae)

The interaction of mating system and nutrient limitation in determining seed production was investigated in the annual, self-compatible plant Lupinus texensis (Fabaceae). Abortion of developing seeds is a major factor limiting seed production in natural populations (17-28%). Selfing rates are generally low (0.02-0.21), suggesting that deleterious recessive genes may be maintained at significant levels in natural populations. The average inbreeding depression associated with seed development is δ = 0.24. Nutrient limitation reduced seed output across experimental treatments by a factor of 0.22 through decreased production of inflorescences, flowers, and ovules, and by a factor of 0.29 through increased abortion of fruits and of seeds within fruits. Competition for resources among fruits increased the frequency of seed abortion. Moreover, a greater proportion of selfed seeds were aborted as the overall abortion rate increased. Estimates of genetic load may therefore only be appropriate if undertaken in the field, and inbreeding depression may vary from year to year simply due to changes in environmental conditions rather than to underlying genetic changes in populations. The existence of inbreeding depression and the high frequency of abortions suggest that selective abortion favoring outcrossed progeny occurs in natural populations of L. texensis.