Life History Evolution in Amphicarpic Plants

Abstract Plants with dimorphic flowers or seeds provide excellent material for the study of life history evolution because the dimorphism often involves measurable differences in morphology, size, number, or genetic relatedness. For amphicarpic plants, the proportion of aerial: subterranean morphs produced is highly variable (from 0 to > 100) and related to both environmental and genetic factors. Plants from aerial seeds produce lower ratios of aerial: subterranean morphs than those from subterranean seeds. Despite substantial variation, subterranean seeds are generally heavier than aerial seeds (but fewer) and produce vigorous seedlings with high survivorship and high fitness. Adaptive advantages of subterranean seeds include retention of offspring in favorable parental microhabitats, protection of seeds from herbivory, predation, or fire, and avoidance of desiccating conditions on the soil surface; potential disadvantages include lack of gene exchange, high energy costs, limited dispersal, and sibling competition. For the few species studied, aerial reproduction is more plastic than subterranean reproduction and more likely to be affected by environmental conditions. Quantitative genetic analyses of a population of the annual grass Amphicarpum purshii have revealed lower heritabilites for subterranean relative to aerial reproductive traits. Subterranean seed number and mass show genetic correlations with shoot mass while aerial seed number and mass do not; seed set percentages of both seed types as well as percentage allocation to both reproductive morphs show negative genetic correlations with shoot mass. In this Amphicarpum population, directional selection on shoot mass may indirectly select for increased subterranean (but not aerial) seed output.     

INFLUENCE OF ENVIRONMENT AND POPULATION ORIGIN ON SURVIVORSHIP AND REPRODUCTION IN RECIPROCAL TRANSPLANTS OF AMPHICARPIC PEANUTGRASS (AMPHICARPUM PURSHII)

Reciprocal transplants of both seeds and seedlings were utilized to determine whether populations of the annual grass Amphicarpum purshii have become locally adapted to specific habitats due to the consistent production of cleistogamous subterranean seeds from year to year. The hypothesis was that subterranean seeds placed in the same habitat as the parents will produce seedlings of greater vigor and adults of higher reproductive capacity than plants from seeds transplanted to a different habitat far removed from the parents. For both seed and seedling transplant experiments involving three sites in the Pine Barrens of New Jersey, the effects of site on shoot dry weight and production of aerial spikelets, subterranean spikelets, and seeds were generally much more significant than the effects of population origin. With one exception, there was no tendency for seedlings (or plants from seeds) replanted into their home sites to outperform alien seedlings (or plants from seeds) transplanted into these same sites. The overriding importance of environmental factors (relative to genetic differences among populations) in determining the phenotypic expression of life history characters, and selection occurring during succession at a site may retard the evolution of genetic adaptation to local habitat conditions in this species.     

The role of seed depth,litter, and fire in the seedling establishment of amphicarpic peanutgrass (Amphicarpum purshii)

Summary Amphicarpum purshii is an annual grass which mostly grows in disturbed areas of the New Jersey Pine Barrens, USA. It is amphicarpic, producing spikelets (and seeds) both above and below the soil surface. Previous research has shown that subterranean seed production ensures reproduction in the event of a major disturbance such as fire and results in rapid post-burn colonization of these sandy habitats. The effects of fire, litter, and seed depth were further examined by planting subterranean seeds at four depths in 16 litter-covered flats buried at ground level and comparing plants arising from burned flats with those in undisturbed litter-covered flats. At 0 and 1 cm depth, rates of seedling emergence were lowest in burned flats. Surface-sown seeds produced seedlings more likely to desiccate. Sowing depth had a greater influence on most measured characters than burning treatments. The mean depth of subterranean seed placement by Amphicarpum is 3.5 cm and this coincides with the seed depth from which plants showed the greatest height growth, shoot biomass, and reproductive output. In a second experiment, subterranean seeds on the bare soil surface in clay pots were more likely to lose viability and less likely to germinate than seeds protected by litter or burial in soil. In addition to providing protection from fire, placement of seeds below ground in the sandy habitat of peanutgrass provides conditions more suitable for seed survival and subsequent seedling establishment.     

RESOURCE ALLOCATION AND REPRODUCTION IN POPULATIONS OF AMPHICARPUM PURSHII (GRAMINEAE)

Five New Jersey populations of Amphicarpum purshii (an annual panicoid grass) were investigated as to total allocation of biomass to reproduction and its distribution between aerial chasmogamous and subterranean cleistogamous inflorescences. Germination of both types of seed was documented, for the first time, in not only the field but under three laboratory temperature regimes. Overall, approximately 29 % of the shoot biomass was allocated to reproduction. On a population basis, subterranean inflorescences accounted for 37–100 % of the reproductive biomass, and these seed (5 × heavier but fewer in number) were the source of most surviving seedlings. Subterranean spikelets contained caryopses significantly more frequently than aerial spikelets. The ratio of the number of viable aerial seed to the number of viable subterranean seed increased from 0/4 to 4/2 with recentness and/or frequency of disturbance. As a pioneer species in secondary succession, Amphicarpum purshii produces a larger subterranean propagule, with greater seedling vigor and a higher probability of local reproductive success, and also a smaller aerial propagule in larger numbers with potentially greater genetic variability.     

Consequences of a mixed reproductive system in the hog peanut,Amphicarpaea bracteata, (Fabaceae)

Summary The forest annual, Amphicarpaea bracteata L. can reproduce via aerial chasmogamous, aerial cleistogamous, and subterranean cleistogamous flowers. Both plant size and light intensity influenced the utilization of the three modes of reproduction. chasmogamous and aerial cleistogamous flower number and the ratio of chasmogamous flowers to the total number of aerial flowers increased with plant size. The latter demonstrated a shift to xenogamy and outbreeding in larger plants. Light intensity indirectly influenced reproductive modes through its infuence on plant size. Seed set by both types of aerial flowers was low and unrelated to plant size. Subterranean seed number and the total dry weight of subterranean seeds per plant increased with size. The subterranean seeds of Amphicarpaea bracteata are thirty-four times larger than the aerial seeds (fresh weight). Under field conditions, subterranean seeds had greater germination after one year than acrial seeds. The plants arising from subterranean seeds were significantly larger and more fecund than those from aerial seeds. Seeds produced by aerial cleistogamous, hand selfpollinated chasmogamous, and naturally pollinated chasmogamous flowers had equivalent germination rates and produced plants of equal size and fecundity. This suggests that the outbred progeny from chasmogamous flowers have no advantage over the inbred progeny from aerial cleistogamous flowers.     

Fruit dimorphism inPhaseolus sublobatus (Fabaceae) and its evolutionary significance

Populations of the annualPhaseolus sublobatus from different ecogeographical zones are genetically differentiated. In twelve populations from the western ghats (Maharashtra range) chasmogamous flowers are arranged in peduncled capitate racemes borne in leaf axils of higher nodes. One population (Poona Race S₄) has additional inflorescences, also with chasmogamous flowers, on the main axis between the cotyledonary node and the ground. When the pods of these flowers ripen, the inflorescence gradually coils and, eventually, gets bury the fruits in the soil. This phenomenon is not known in any other plant.—The seed-coat patterns as revealed by SEM, and germination behaviour of both aerial and subterranean seeds are similar, and both types of seeds lack dormancy.—Heavy seed predation selection pressure is probably the principal cause for the evolution of fruit dimorphism inP. sublobatus. The origin of geocarpy in relation to fruit dimorphism and seed predation is discussed.     

Reproductive versatility in legumes: the case of amphicarpy in Trifolium polymorphum

Amphicarpy is a fascinating reproductive strategy, defined as fruit produced both below the soil surface and as aerial fruit on the same plant. Trifolium polymorphum is a grassland species subject to herbivory that combines amphicarpy with vegetative reproduction through stolons. Underground flowers have been described as obligate autogamous and aerial ones as self‐compatible allogamous, with aerial floral traits favouring cross‐pollination. In the present work we performed different pollination treatments on aerial flowers to analyse rates of pollen tube development and offspring fitness, measured as fruit set, seed production and germination percentage. This last variable was compared to that of seeds produced underground. No significant differences were found between fruit set in self‐ and cross‐pollinations. Seed production was higher in self‐pollinations, which is consistent with the higher rate of pollen tube development observed in self‐crosses. Spontaneous self‐pollination is limited in aerial flowers; thus pollen transfer by means of a vector is required even within the same flower. Germination tests showed that aerial seeds produced after self‐ and cross‐pollination did not differ in fitness, but underground seeds had higher germination percentage than aerial ones. Thus, we conclude that T. polymorphum has a mixed mating system. In grasslands with heavy grazing pressure, clonal propagation and underground seed production ensure persistence in the field. An intermediate level of selfing in aerial flowers ensures offspring, but morphological (herkogamy) and functional (dicogamy) floral traits maintain a window to incorporate genetic variability, allowing the species to tolerate temporal and spatial pressures.     

Amphicarpum purshii and the “pessimistic strategy” in amphicarpic annuals with subterranean fruit

Summary Amphicarpum purshii Kunth is an annual amphicarpic grass of recently disturbed sandy areas in the Coastal Plain of eastern North America, producing small aerial and larger subterranean seeds. At study sites on the eastern edge of the Pine Barrens of New Jersey, the temporal aspects of reproductive allocation were investigated through biweekly whole-plant field harvests of 25 plants each throughout the 1980 growing season, and the survivorship of plants arising from aerial and subterranean seeds was monitored in a total of 10 high and 10 low density quadrats at two sites differeing in soil moisture. Plants arising from subterranean propagules began to allocate early (July 16) considerable energy to subterranean reproduction (ca 40% of whole-plant dry weight by the last half of August); allocation to aerial culms and flowers did not begin until mid-August and never exceeded 3% on a population basis. Plants arising from aerial propagules (aerial plants) produced only subterranean flowers, and these flowers first appeared 2 wk later than they did on subterranean plants. Survivorship was greater for the subterranean seedlings at both the dry and wet sites and at low and high densities, and aerial plants showed significantly less total growth and seed production than subterranean plants. The pessimistic strategy (early allocation of energy to large subterranean propagules) in the annual Amphicarpum has its selective basis in the relative vigor, survival, and timing and amount of reproduction of the two types of seedlings, and appears comparable to the allocation strategies of at least eight other amphicarpic annuals in at least five other families of plants.     

GENETIC AND ENVIRONMENTAL VARIATION IN LIFE-HISTORY TRAITS OF A MONOCARPIC PERENNIAL: A DECADE-LONG FIELD EXPERIMENT

Directional and stabilizing selection tend to deplete additive genetic variance. On the other hand, genetic variance in traits related to fitness could be retained through polygenic mutation, spatially varying selection, genotype-environment interaction, or antagonistic pleiotropy. Most estimates of genetic variance in fitness-related traits have come from laboratory studies, with few estimates of heritability made under natural conditions, particularly for longer lived organisms. Here I estimated additive genetic variance in life-history characters of a monocarpic herb, Ipomopsis aggregata, that lives for up to a decade. Experimental crosses yielded 229 full-sibships nested within 32 paternal half-sibships. More than 5000 offspring were planted as seeds into natural field sites and were followed in most cases through their entire life cycle. Survival showed substantial additive genetic variance (genetic coefficient of variation ≈ 54%). Small differences at seedling emergence were magnified over time, such that the genetic variability in survival was only detectable by tracking the success of offspring for several years starting from seed. In contrast to survival, reproductive traits such as flower number, seeds per flower, and age at flowering showed little or no genetic variability. Despite relatively high levels of additive genetic variation for some life-history characters, high environmental variance in survival resulted in very low heritabilities (0–9%) for all of these characters. Maternal effects were evident in seed mass and remained strong throughout the lengthy vegetative period. No negative genetic correlations between major components of female fitness were detected. Mean corolla width for a paternal family was, however, negatively correlated with the finite rate of increase based on female fitness. That negative correlation could help to maintain additive genetic variance in the face of strong selection through male function for wide corollas.     

Genetic variation and self-incompatibility within and outside a Rorippa hybrid zone (Brassicaceae)

Rorippa amphibia and R. sylvestris are both self-incompatible, perennial species. In northern Germany they show a geographic pattern of interspecific gene flow: R. amphibia and R. sylvestris form a hybrid zone at the river Elbe but do not hybridise elsewhere in northern Germany. In the present paper we analyse the relationship between molecular diversity, seed set and seed germination within and outside the hybrid zone. In both species we observed high genetic variation and high seed set in the hybrid zone, and low genetic variation and low seed set outside the hybrid zone. In R. amphibia, high genetic variation is correlated with high seed set, but with low germination rates of the seeds. The results of an isolation experiment show that self-incompatibility is maintained in the parental species and their hybrids. The impact of genetic self-incompatibility on hybrid zone formation and hybrid fitness is discussed.     

ENVIRONMENTAL AND GENETIC MATERNAL EFFECTS ON SEED CHARACTERS IN NEMOPHILA MENZIESII

Nuclear genetic, maternal genetic and maternal environmental effects on seed characters were estimated in the California native annual plant Nemophila menziesii using two greenhouse crosses. In one cross, according to a nested mating design, the narrow sense heritability of seed weight was small (3.9%). A subset of full-sib progenies produced in this cross was grown singly and in competition with the introduced grass Bromus diandrus. In a second cross, these plants were used as mothers (dams) and were each mated to the same three sires. Seeds produced by mothers competing with B. diandrus showed a significant reduction in weight, increase in time to germination, and increase in the incidence of dormancy, when compared to seeds from mothers grown singly. Significant sire components were found for time to germination and incidence of dormancy. Maternal genetic variation for seed weight was largely expressed as maternal genotype by maternal environment interaction, and showed no significant maternal genetic main effect. Time to germination and dormant fraction showed a relatively large maternal genetic effect. Evolution of seed characters in N. menziesii is more likely to occur via indirect response to selection among maternal plants than among the seeds themselves. Maternal genotype by maternal environment interaction could potentially contribute to the maintenance of maternal genetic variation in seed weight, but this does not appear likely for dormancy.     

DISPERSAL OF HETEROMORPHIC SEEDS IN AMPHICARPAEA BRACTEATA (FABACEAE)

The dispersal of the chasmogamous, aerial cleistogamous, and subterranean cleistogamous seeds of Amphicarpaea bracteata was examined. The chasmogamous and aerial cleistogamous seeds are ballistically dispersed. Chasmogamous seeds were dispersed farther than the aerial cleistogamous seeds due to the height advantage of the chasmogamous pods. There was no difference in the firing angle or the initial velocity of the seeds discharged from the two aerial pod types. The subterranean cleistogamous seeds are “dispersed” the shortest distance by the elongation of runners. Differential dispersal of the three types of seeds may be a factor involved in the evolution of cleistogamy in A. bracteata.     

Fitness components and the determinants of fecundity in populations of a native perennial grass (Tridens flavus)

Understanding intraspecific variation in traits that determine fitness is foundational to a trait-based approach to plant ecology. This study examined fitness components during 3 years of reproduction in a polycarpic perennial bunchgrass (Tridens flavus) native to eastern North America that could prove useful in revegetating disturbed habitats. Plants were cultured from seeds of five populations in central New Jersey, USA, and planted in July 2015 into two undisturbed gardens 30 m apart that differed in availability of sunlight and soil moisture. Following flowering in 2016, 2017 and 2018, the number of panicles, seed set, seed number (fecundity) and seed mass were recorded. Final dry aboveground mass was determined. Seed set was high (>70%) in all populations and gardens. Panicle production varied with population and was strongly correlated with fecundity, but populations were not differentiated for other fitness components. Panicle and seed number were greatest in the drier garden with greater daily light availability. Mass per seed was reduced as more seeds were produced in the second and third year but showed low variation compared to fecundity. Vegetative mass was the most important variable determining fecundity. Close proximity of sampled sites and an outcrossed, wind-pollinated mating strategy may have precluded detection of differentiation among T. flavus populations in the common gardens. High seed set, prodigious seed production on multiple panicles and high seed germinability and overwinter survival account for the occurrence of large populations of this native grass along roadsides and within successional fields and young woodlands throughout the region.     

茶条槭自然种群种子和果实表型多样性研究

为探析茶条槭(Acer ginnala)种子和果实表型差异程度及变异规律,该研究采用巢式方差分析、主成分分析、聚类分析等方法,对茶条槭主要分布区内7个种群种子和果实的12个表型性状进行比较研究,分析茶条槭种群间和种群内的表型多样性以及与地理生态因子的关系。结果表明:除种子长/宽(SLW)外,其它11个表型性状在茶条槭种群间和种群内均具有显著差异;各性状平均变异系数(CV)为13.90%,变异幅度为8.14%～32.08%;种群间翅果性状的平均变异系数(15.63%)高于种子性状的平均变异系数(8.71%),表明种子性状的稳定性高于果实性状。主成份分析中,果实形态特征对茶条槭种群表型的贡献率大于种子贡献率。种群间表型分化系数为35.47%,说明种群内变异(64.53%)大于种群间变异(35.47%),暗示变异主要来源于种群内变异。茶条槭种子和果实表型性状受地理生态因子影响较小,主要受自身遗传因素的影响。基于种群间欧式距离的聚类分析将茶条槭7个种群分为2大支,没有严格按照地理距离而聚类,表明茶条槭种群表型性状变异的不连续性。茶条槭不同种群种子和果实具有较高的表型多样性,与其分布范围和生物学特性有关。     

黄土丘陵沟壑区主要物种植冠种子库动态及其生态策略

植冠种子库是植物适应环境并应对外界干扰的种子生态策略之一,研究了黄土丘陵沟壑区12种主要植物植冠种子库动态,结果表明:杠柳不具有植冠种子库,其他11种植物均具有植冠种子库;除了黄刺玫种子在翌年5月达到脱落高峰,其他植物大部分种子在冬季脱落,其中杠柳、达乌里胡枝子、茭蒿、黄柏刺和水栒子的大部分种子脱落集中偏早,铁杆蒿和土庄绣线菊的大部分种子脱落集中偏晚;植冠宿存对大部分植物种子的萌发特性表现为促进作用;但不同植物种子的萌发时滞对植冠宿存响应差异较大;9种植物种子在植冠上宿存至翌年2月底,其种子活力仍能维持达60%以上;该区植物表现出不同的植冠种子库策略,通过不同的方式来减少干扰的威胁,提高成功萌发与更新的几率,它们或具有较大规模的宿存量、或调控种子萌发特性、或提高种子维持活力的百分比。此外,全面了解该区植物形成植冠种子库的机理及对应的生态策略还有待于全面、深入的研究。     

Offspring fitness in relation to population size and genetic variation in the rare perennial plant species Gentiana pneumonanthe (Gentianaceae)

Seeds were sampled from 19 populations of the rare Gentiana pneumonanthe, ranging in size from 5 to more than 50,000 flowering plants. An analysis was made of variation in a number of life-history characters in relation to population size and offspring heterozygosity (based on seven polymorphic isozyme loci). Life-his-tory characters included seed weight, germination rate, proportion of seeds germinating, seedling mortality, seedling weight, adult weight, flower production per plant and proportion of plants flowering per family. Principal component analysis (PCA) reduced the dataset to three main fitness components. The first component was highly correlated with adult weight and flowering performance, the second with germination performance and the third component with seed and seedling weight and seedling mortality. The latter two components were considered as being maternally influenced, since these comprised life-history traits that were significantly correlated with seed weight. Multiple regression analysis showed that variation in the first fitness component was mainly associated with heterozygosity and not with population size, while the third fitness component was only correlated with population size and not with heterozygosity. The latter relationship appeared to be non-linear, which suggests a stronger loss of fitness in the smallest populations. The second (germination) component was neither correlated with population size nor with genetic variation. There was only a weak association between population size, heterozygosity and the population coefficients of variation for each life history character. Most correlation coefficients were negative, however, which suggests that there is more variation among progeny from smaller populations. We conclude that progeny from small populations of Gentiana pneumonanthe show reduced fitness and may be phenotypically more variable. One of the possible causes of the loss of fitness is a combination of unfavourable environmental circumstances for maternal plants in small populations and increased inbreeding. The higher phenotypic variation in small populations may also be a result of inbreeding, which can lead to deviation of individuals from the average phenotype through a loss of developmental stability.     

贵州省野生皂荚种实表型性状多样性分析

为深入了解贵州省野生皂荚(Gleditsia sinensis)荚果表型性状的遗传多样性及其变异类型,为皂荚的遗传改良、种质鉴定、亲本选择以及品种培育奠定理论基础。该研究以贵州省7个野生皂荚群体70个个体为研究对象,采用方差分析、主成分分析、相关性分析及多性状综合评价等方法对皂荚群体的10个种实表型性状进行系统分析和综合评价。结果显示：(1)所测皂荚的表型性状差异在群体内均达到极显著水平(P＜0.01);在群体间,除每荚粒数、种子宽、种子长宽积以及种子长宽比之外,其余表型性状的差异均达极显著水平(P＜0.01)。(2)7个居群野生皂荚各性状平均变异系数为21.16%,其中凯里市(P4)居群的变异系数最高(24.44%);居群间荚果的变异(29.22%)高于种子的变异(11.04%),且变异主要来自于群体内。(3)相关分析显示,皂荚种实各性状之间存在不同程度的关联性;主成分分析显示,前4个主成分(皂荚种子大小、单个荚果出籽数量、种子形态指数因子、与荚果长和种子厚相关的因子)的累积贡献率达69.783%,可基本反映皂荚表型性状的大部分信息;以10个种实性状对皂荚野生群体进行综合评价发现,来自于惠水县(P7)群体的皂荚种实性状综合评价最高。研究表明,贵州省野生皂荚在群体间及群体内具有丰富的表型变异,且群体内的变异大于群体间的变异,变异主要来自于群体内。     

REPRODUCTIVE BEHAVIOR OF AMPHICARPAEA BRACTEATA (LEGUMINOSAE), AN AMPHICARPIC ANNUAL

Amphicarpaea bracteata, an annual legume common in woodland communities in the eastern United States, produces three distinct types of flower: subterranean cleistogamous (SCL), aerial cleistogamous (ACL), and aerial chasmogamous (ACH). We sought to quantitatively describe the growth and reproduction of four diverse A. bracteata populations in an effort to explain the adaptive significance of this species’ variable reproductive modes. Virtually all plants develop 1 to 3 SCL seeds on subterranean cotyledonary shoots, starting in late July. Larger plants produce additional SCL seeds later from the tips of axillary shoots which bury. Seeds produced by SCL flowers are large (39 to 134 mg), have restricted dispersal, lack dormancy, have high germination , and produce relatively vigorous seedlings. ACL flowers develop from early August through plant senescence in early October, while ACH flowers appear in a pulse in late August. The number of each aerial flower type was positively correlated with plant size. The ACL flowers take significantly less time to develop mature fruits than do ACH flowers. Both aerial seeds are smaller than the subterranean (mean weights: ACL 12.1, ACH 8.7 mg), have lower germination (ACL 20.2%, ACH 15.3%), appear relatively resistant to environmental extremes, and have the potential for more widespread dispersal. Plants derived from subterranean seeds are much larger than plants derived from aerial seeds, which makes them more likely to produce axillary SCL and aerial seeds. Only larger plants derived from subterranean seeds and growing in favored sites produce ACH flowers. This imposes an alternation of selfing with outcrossing generations. This pattern of reproductive behavior in A. bracteata is similar to that found in several other amphicarpic species.     

Population differentiation in the tiller architecture of Microstegium vimineum (Poaceae) in relation to habitat

Grass tillers grow by addition of modular units known as phytomers. Differences in phytomer organ size produce subindividual variation with potential adaptive value. Here, patterns in organ mass along tillers in the invasive annual Microstegium vimineum are related to habitat and tiller architecture. In an earlier study, seed families were collected from two populations: one from a sunny, woodland edge and the other from a shady understory in New Jersey, USA. Plants from these seeds were grown in a greenhouse. Phytomers along primary tillers were divided into culms, leaves, and cleistogamous or chasmogamous spikelets and seeds, dried and weighed. These data were used to examine the quantitative genetics of subindividual variation among families and populations, and the relationship of tiller fitness (based on total seed mass) to the mean or subindividual variance of phytomer traits. Phytomer position along a tiller was the major determinant of organ mass. Leaf mass increased from basal to upper nodes; cleistogamous reproductive mass decreased from upper to lower nodes. Phytomer organs were heaviest in the population from the sunny habitat. Family explained < 18% of variation in organ mass. Tiller fitness was positively correlated with mean culm, and leaf mass, but negatively correlated with coefficients of variation. Field‐collected tillers showed evidence of selection for increased leaf mass. Subindividual variation in M. vimineum is mostly due to phytomer position along a tiller and the prevailing light environment. Differentiation between sunny and shady populations suggests selection favors heavier phytomer leaves and culms, especially in the shady understory where this species is most invasive.     

高山松种实性状与生殖适应性

高山松(Pinus densata)是云南松(P. yunnanensis)与油松(P. tabulaeformis)自然杂交产生的二倍体杂种,是植物同倍体杂交物种形成的典型案例。高山松分布于青藏高原东南缘,占据了两个亲本种都不能正常生长的高海拔地带。良好生殖适应性的产生是物种形成的关键,但对高山松在高海拔生境下生殖适应性的变异与杂交起源遗传背景关系的研究尚未见报道。该研究通过对来自该种分布区内6个代表居群样本、13个反映高山松生殖适应性的种实性状的分析,探讨了其生殖性状在居群水平的变异模式,揭示了具有不同遗传组成和起源历史的高山松居群的生殖特性。研究结果表明:6个研究居群在种实性状上存在显著差异;可育种鳞数、总种鳞数、总种子数、可育种鳞率和可育种鳞密度等与纬度呈显著负相关关系;球果结种率与经度和生态梯度值呈显著的正相关关系;胚珠败育率则与经度和生态梯度值呈显著的负相关关系;球果长度也呈现了与经度的显著负相关。高山松球果结种率很高,平均为74%;球果的结种数、种子长和种翅长与云南松、油松及其它松属双维管亚属多数种接近;球果长度、球果总种鳞数接近油松的报道。这些结果表明,高山松在亲本种不能正常生长、繁衍的高原环境下具备正常的生殖能力;其种实特征在居群间分化显著并呈现一定的地理梯度变化。居群间的显著差异反映了其杂种起源特性,与高山松居群的遗传背景有关,也反映了处在异质生态环境中的高山松居群具有各自的生殖生态进化趋向。