放牧对小嵩草草甸生物量及不同植物类群生长率和补偿效应的影响

基于小嵩草(Kobresia parva)草甸连续2 a的牦牛放牧试验,研究了暖季和冷季放牧草场地上地下生物量及其分配规律、不同植物类群的绝对生长率生长率,探讨了放牧制度和放牧强度对不同植物类群补偿效应的影响。结果表明,随着放牧强度的增加地上总生物量呈减小趋势,放牧强度对暖季草场地上总生物量的影响极显著（P?0.01）,对冷季草场地上总生物量的影响不显著（P?0.05）;两季放牧草场各土壤层地下生物量随放牧强度的增加呈明显下降趋势,放牧强度对暖季放牧各土壤层地下生物量的影响显著（P?0.05）,对冷季放牧各土壤层地下生物量的影响不显著（P?0.05）;冷季放牧草场牧草生长季地下生物量与地上生物量的比值随放牧强度的增大而减小,暖季放牧草场对照区地下生物量与地上生物量的比值低于轻度放牧和中度放牧、高于重度放牧;暖季放牧草场各放牧处理不同植物类群均存在超补偿生长,但莎草科和禾本科植物的超补偿生长在8月份,阔叶植物的超补偿生长发生在6月和7月份,禾本科植物的超补偿生长效应强于莎草科植物和阔叶植物,轻度和中度放牧的补偿效应更明显;冷季放牧下不同植物类群也存在超补偿生长,但补偿效应不明现。因此,暖季适度（轻、中度）放牧利用更有利于产生超补偿生长,而重度利用对植被的稳定产生潜在的不利影响。     

Complex spatial clonal structure in the macroalgae Fucus radicans with both sexual and asexual recruitment

In dioecious species with both sexual and asexual reproduction, the spatial distribution of individual clones affects the potential for sexual reproduction and local adaptation. The seaweed Fucus radicans, endemic to the Baltic Sea, has separate sexes, but new attached thalli may also form asexually. We mapped the spatial distribution of clones (multilocus genotypes, MLGs) over macrogeographic (>500 km) and microgeographic (<100 m) scales in the Baltic Sea to assess the relationship between clonal spatial structure, sexual recruitment, and the potential for natural selection. Sexual recruitment was predominant in some areas, while in others asexual recruitment dominated. Where clones of both sexes were locally intermingled, sexual recruitment was nevertheless low. In some highly clonal populations, the sex ratio was strongly skewed due to dominance of one or a few clones of the same sex. The two largest clones (one female and one male) were distributed over 100–550 km of coast and accompanied by small and local MLGs formed by somatic mutations and differing by 1–2 mutations from the large clones. Rare sexual events, occasional long‐distance migration, and somatic mutations contribute new genotypic variation potentially available to natural selection. However, dominance of a few very large (and presumably old) clones over extensive spatial and temporal scales suggested that either these have superior traits or natural selection has only been marginally involved in the structuring of genotypes.     

Genetic diversity of the expansive grass Brachypodium pinnatum in a changing landscape: Effect of habitat age

Perennial grasses constitute a major group of species showing a dramatic decline of biodiversity in successional plant communities. Using AFLP markers, we examined 12 populations of the expansive grass Brachypodium pinnatum differing in habitat age (30–50, ca. 100 and >300 years old) in order to determine whether clonal diversity of populations, genetic variation, and the relative importance of clonal propagation versus sexual reproduction change with grassland age. Five AFLP primer combinations gave a total of 517 bands, 79% of which were polymorphic. 314 different multilocus lineages were distinguished among the 453 samples analyzed. The number of genotypes (G) and clonal richness (R) decreased with habitat age, while the distribution of the frequency of genets changed from many clones of similar size to dominance by one or a few large clones. We consider these results to give evidence of significant role of sexual reproduction in the early phases of colonization and prevalence of clonal growth and competitive exclusion of less adapted genotypes in the later ones. However, habitat age had only marginal effect on genetic diversity, as percentage of polymorphic loci (PPL) within all the populations analyzed was similar, viz. 38.6–43.5%.     

Testing for clonal propagation

The conceptual basis for testing clonal propagation is reconsidered with the result that two steps need to be distinguished clearly: (1) specification of the characteristics of multilocus genotype frequencies that result from sexual reproduction together with the kinds of deviations from these characteristics that are produced by clonal propagation, and (2) a statistical method for detecting these deviations in random samples. It is pointed out that a meaningful characterization of sexual reproduction reflects the association of genes in (multilocus) genotypes within the bounds set by the underlying gene frequencies. An appropriate measure of relative gene association is developed which is equivalent to a multilocus generalization of the standardized gametic disequilibrium (linkage disequilibrium). Its application to the characterization of sexually produced multilocus genotypes is demonstrated. The resulting hypothesis on the frequency of a sexually produced genotype is tested with the help of the (significance) probability of obtaining at least two copies of the genotype in question in a random sample of a given size. If at least two copies of the genotype are observed in a sample, and if the probability is significant, then the hypothesis of sexual reproduction is rejected in favor of the assumption that all copies of the genotype belong to the same clone. Common testing approaches rest on the hypothesis of completely independent association of genes in genotypes and on the (significance) probability of obtaining at least as many copies of a genotype as observed in a sample. The validity of these approaches is discussed in relation to the above considerations and recommendations are set out for conducting appropriate tests.     

高寒草甸矮蒿草种群繁殖对策的研究

繁殖对策是指生物对环境的生殖适应趋势 ,是资源或能量向生存、生长和生殖等活动中最适分配的结果 ,在不同的环境中具有其独特的表现形式。研究植物在不同环境中的繁殖对策可以反映出植物对环境的适应能力和在该生境中的生殖潜能。国内外学者对植物繁殖对策的研究已有不少报道[2 ,4 ,5,6] 。但对高寒草甸矮嵩草 (Ｋｏｂｒｅｓｉａｈｕｍｉｌｉｓ)种群繁殖对策的研究报道甚少。矮嵩草是青藏高原矮嵩草草甸的建群种 ,它具有草质柔软、营养丰富、热值含量较高等特点 ,是青藏高原重要的可更新草地资源。本研究对矮嵩草的繁殖对策进行了较全面、…     

Quantifying the levels of sexual reproduction and clonal spread in the invasive plant, Rosa multiflora

Rosa multiflora Thunb., (Rosaceae), an invasive plant in the eastern U.S., was introduced into the U.S. in the early 1800s and was widely planted in the 1940s as a living fence, for wildlife cover, and to prevent soil erosion. This species spread rapidly from these original plantings via seed dispersal (sexual reproduction) and clonal spread, invading pasture and wooded areas. In this study we used allozyme markers to test for significant differences in the levels of asexual and sexual spread in large (>9 m cirumference) verses small (<2 m circumference) patches of R. multiflora and in pasture verses park settings. Although larger patches of R. multiflora tended to be dominated by one genotype, they exhibited significantly greater genetic diversity and inputs from sexual reproduction than did small patches; all large patches (N = 10) contained multiple unique genotypes. In contrast six of ten smaller patches of R. multiflora, consisted of a single genotype, though three patches had two genotypes and one had three unique genotypes. Similar analyses revealed clonal structure in R. multiflora populations both park and pasture habitats but with significantly greater genetic diversity and sexual inputs in the former than the latter. These results are consistent with a model of invasive spread involving clonal spread, sexual reproduction, and bird-mediated seed dispersal into established patches. Sexual inputs appear to be highest in larger patches and park habitats where perching sites for birds are most abundant. This flexible reproduction system likely contributes to the invasiveness of R. multiflora and to current management failures.     

高寒草甸放牧扰动与两种植物的反应研究

对不同放牧强度下矮嵩草草甸植物垂穗披碱草和鹅绒委陵菜的生长和繁殖特性进行了5年研究。结果表明,垂穗披碱草株高、分蘖数、地上物量、地下生物量、有性繁殖效力在不放牧或轻度放牧下最大,营养生长效力在重度放牧中最大,而贮藏效力在重度放牧中最小。鹅绒委陵菜叶数、匍匐茎长度、地上生物量、匍匐茎生物量、开花数、生殖生物量、有性繁殖效力及营养繁殖效力在放牧区显著高于不放牧的对照区。说明适当的放牧干扰有利于垂穗披碱草营养生长,但放牧过重会抑制其有性繁殖;由于放牧采食减少了种间竞争影响,从而提高了鹅绒委陵菜营养,生长效力和有性繁殖效力。两物种地下贮藏物的比例随着放牧强度增加显著减少。放牧强度对高寒草甸的群落演替方向有显著影响。     

Clonal diversity in the rare Calamagrostis porteri ssp. insperata (Poaceae): comparative results for allozymes and random amplified polymorphic DNA (RAPD) and intersimple sequence repeat (ISSR) markers

Four populations of the rare, highly clonal grass Calamagrostis porteri ssp. insperata were examined using allozymes and the two polymerase chain reaction (PCR)-based markers, random amplified polymorphic DNA (RAPD) and intersimple sequence repeat (ISSR) bands. Only one of the 15 allozyme loci was variable and two alleles were detected, both of which were found in two populations, while only one genotype was detected in the other two populations. ISSR and RAPD markers detected more genotypes within populations than did allozymes. ISSR markers detected more diversity than RAPD markers in three of the four populations examined. In one population, no RAPD diversity was found whereas eight different genotypes were found among the 10 plants with ISSR markers. This diversity is present despite rare flowering, no documented occurrence of seed set in natural populations and very low seed set with experimental pollinations, all of which suggest that sexual reproduction rarely occurs. The subspecies is self-compatible, but seed initiation is lower in selfed ovules; also, there is high embryo abortion regardless of pollen source. Variation detected by RAPD and ISSR primers may reflect higher levels of sexual reproduction in the past, very rare sexual reproduction in extant populations, somatic mutations, or a combination of the three. Although the PCR-based markers identify several multilocus genotypes within populations, it is not known whether these all represent distinct genets generated by sexual reproduction or result from somatic mutations in the old, perennial and highly clonal plants.     

Prolonged clonal growth: escape route or route to extinction?

Many plant species have the capability to reproduce sexually as well as clonally. The balance between clonal reproduction and sexual reproduction varies between different species. It was estimated that 66.5% of all central European flora may form independent but genetically identical daughter plants. Also within species there is great variation in the ratio clonal/sexual reproduction. Clonal reproduction can be considered as an alternative life cycle loop that allows persistence of a species in the absence of the ability to complete the normal life cycle (i.e. seed production, germination and recruitment). Plant populations exhibiting prolonged clonal growth have been referred to as 'remnant populations'. A remnant population in general is defined as "a population capable of persistence during extended time periods despite a negative population growth rate (λ<1) due to longlived life stages and life cycles, including loops, that allow population persistence without completion of the whole life cycle". Here we argue that prolonged and nearly exclusive clonal growth through environmental suppression of sexual reproduction can ultimately lead to local sexual extinction and to monoclonal populations of a species, and that this may imply significant consequences for population viability. Especially obligate or mainly outcrossing clonal plant species may be vulnerable for sexual extinction. We argue that the consequences of reduced sexual recruitment in clonally propagating plants may be understudied and underestimated and that a re-evaluation of current ideas on clonality may be necessary.     

牦牛放牧强度对小嵩草草甸两季轮牧草场植物群落数量特征的影响

基于小嵩草(Kobresia parva)草甸连续2年的牦牛放牧控制试验,研究了暖季和冷季放牧草场植物群落数量特征的变化.结果表明:在2年的放牧期内,小嵩草草甸2季草场优良牧草的盖度随放牧强度的提高呈降低趋势,而杂草的盖度呈增加趋势;2季草场对照、轻度放牧和中度放牧组的优势种均为小嵩草和垂穗披碱草,但暖季草场重度放牧下主要优势种变为鹅绒委陵菜和阿拉善马先蒿,冷季草场重度放牧下小嵩草和鹅绒委陵菜为主要优势种;放牧强度之间的差异越大,2季草场各放牧处理间群落的相似性程度越低,说明放牧强度是引起群落差异的主要原因,也是群落变化的主导因子.2季草场各放牧处理的物种丰富度、多样性指数、均匀度指数在对照组最低,中度放牧组最高,其指数排序为:对照＜轻度放牧＜重度放牧＜中度放牧,这一结果支持“中度干扰理论”.     

Population genetic consequences of extreme variation in sexual and clonal reproduction in an aquatic plant

Most plants combine sexual reproduction with asexual clonal reproduction in varying degrees, yet the genetic consequences of reproductive variation remain poorly understood. The aquatic plant Butomus umbellatus exhibits striking reproductive variation related to ploidy. Diploids produce abundant viable seed whereas triploids are sexually sterile. Diploids also produce hundreds of tiny clonal bulbils, whereas triploids exhibit only limited clonal multiplication through rhizome fragmentation. We investigated whether this marked difference in reproductive strategy influences the diversity of genotypes within populations and their movement between populations by performing two large-scale population surveys (n = 58 populations) and assaying genotypic variation using random amplified polymorphic DNA (RAPDs). Contrary to expectations, sexually fertile populations did not exhibit higher genotypic diversity than sterile populations. For each cytotype, we detected one very common and widespread genotype. This would only occur with a very low probability (< 10-7) under regular sexual recombination. Compatibility analysis also indicated that the pattern of genotypic variation largely conformed to that expected with predominant clonal reproduction. The potential for recombination in diploids is not realized, possibly because seeds are outcompeted by bulbils for safe sites during establishment. We also failed to find evidence for more extensive movement of fertile than sterile genotypes. Aside from the few widespread genotypes, most were restricted to single populations. Genotypes in fertile populations were very strongly differentiated from those in sterile populations, suggesting that new triploids have not arisen during the colonization of North America. The colonization of North America involves two distinct forms of B. umbellatus that, despite striking reproductive differences, exhibit largely clonal population genetic structures.     

High genotypic diversity and strong spatial structure in populations of Trifolium alpestre with low seed production

Studies on clonal plants indicate that the proportion between clonal and sexual reproduction can be variable, depending on local habitat conditions and the biological characteristics of the species. In the present study, we assessed this question in Trifolium alpestre by assaying genetic diversity and spatial genotypic structure of natural populations with the use of allozyme markers. Populations revealed high genetic diversity as well as strong spatial structure of multilocus genotypes. The values of genetic diversity were moderately high. Spatially aggregated, identical genotypes spread up to 15 m along linear transects and across 4‐m² plots indicate extensive clonal propagation within populations. However, the existence of numerous unique and small‐sized clones reflects significant contribution from sexual reproduction. Spatially and temporarily stochastic soil disturbances have evidently opened new opportunities for the successful sexual recruitment from the permanent soil seed bank and thus counteracted losses of genetic and genotypic diversity. Seed production in all populations during the three study years was low, in average up to 1.5–2.4 seeds per shoot. The almost total lack of seed set for 57 bagged flower heads on genotypes grown in a common garden indicates that T. alpestre needs pollinators for seed production.     

Patterns of clonal diversity in three species of sub-arctic willow (Salix lanata, Salix lapponum and Salix herbacea)

Sub-arctic willow scrub is an endangered habitat in Britain, and typically occurs on steep crags inaccessible to grazing animals. These willows can reproduce both sexually and asexually, although the relative importance of each is unknown. Knowledge of reproductive mode is important for the design of grazing management and restoration programmes. Accordingly, clonality was assessed in the largest stand of sub-arctic willow scrub in the UK, focusing on Salix lanata and S. lapponum. Little evidence of clonal growth was detected; most individuals possessed distinct multi-locus genotypes. Thus despite the capacity for vegetative reproduction, and seedlings being rarely observed, sexual reproduction is the predominant means of perpetuation and dispersal at this site. We also examined clonal growth in a common willow species (Salix herbacea) that occupies a different habitat type (exposed mountain tops and ridges). Multiple individuals shared identical genotypes up to 7 m apart, suggesting an important role for clonal growth in local patch formation in this species.     

草地植物种群繁殖对策研究

植物的繁殖包括有性繁殖和无性繁殖两大类型,克隆繁殖是一种较为特殊的营养繁殖方式。本文综述了草地种子植物的生殖分配及生殖投资,克隆生长以及放牧对草地植物种群繁殖的影响。植物种群生物量、能量和养分生殖分配是植物种群生殖分配的重要内容,不同植物在结实期营养元素及能量的配置上有着显著的区别,这可能是植物在长期进化过程中形成的生殖对策,是适应环境的结果。在种群水平上,中等强度以上的放牧干扰有利于植物的克隆生长,但有性生殖减弱。草原植物发达的营养繁殖或克隆生长方式是对放牧的适应性对策。     

Characterization of CO2 flux in three Kobresia meadows differing in dominant species

Aims Kobresia meadows, the dominant species of which differ in different habitats, cover a large area of alpine grassland on the Qinghai-Tibetan Plateau and act as potential CO₂ sinks. Kobresia meadows with different dominant species may differ in carbon sink strength. We aimed to test the hypothesis and to clarify the differences in CO₂ sink strength among three major Kobresia meadows on the plateau and the mechanisms underlying these differences.Methods We measured the net ecosystem exchange flux (NEE), ecosystem respiration flux (ER), aboveground biomass (AGB) and environmental variables in three Kobresia meadows, dominated by K. pygmaea, K. humilis, or K. tibetica, respectively, in Haibei, Qinghai. NEE and ER were measured by a closed-chamber method. Environmental variables, including photosynthetic photon flux density (PPFD), air and soil temperature and air and soil moisture, were monitored during the above flux measurements.Important findings The measured peak AGB increased with soil water content and was 365, 402 and 434 g dry weight m^{-2<-sup> for K. pygmaea, K. humilis and K. tibetica meadow, respectively. From the maximum ecosystem photosynthetic rate in relation to PPFD measured during the growing season, we estimated gross ecosystem photosynthetic potential (GEP max) as 22.2, 29.9 and 37.8 μmol CO₂ m-2<-sup> s-1 for K. pygmaea, K. humilis and K. tibetica meadow, respectively. We estimated the respective gross primary production (GPP) values as 799, 1-063 and 1?158 g C m-2<-sup> year-1 and ER as 722, 914 and 1-011 g C m-2<-sup> year-1. Average net ecosystem production (NEP) was estimated to be 76.9, 149.4 and 147.6 g C m-2<-sup> year-1 in K. pygmaea, K. humilis and K. tibetica meadows, respectively. The results indicate that (i) the three meadows were CO₂ sinks during the study period and (ii) Kobresia meadows dominated by different species can differ considerably in carbon sink strength even under the same climatic conditions, which suggests the importance of characterizing spatial heterogeneity of carbon dynamics in the future.}     

Small-scale species richness and its spatial variation in an alpine meadow on the Qinghai-Tibet Plateau

We investigated how the high small-scale species richness of an alpine meadow on the Qinghai-Tibet Plateau, China, is maintained. This area is characterized by strong wind and severe cold during long winters. In winter, most livestock is grazed on dead leaves in small pastures near farmers’ residences, whereas in the short summer, livestock is grazed in mountainous areas far from farmers’ residences. The number of plant species and the aboveground biomass were surveyed for three adjacent pastures differing in grazing management: a late-winter grazing pasture grazed moderately from 1 February to 30 April, an early-winter grazing pasture grazed lightly from 20 September to late October, and a whole-year grazing pasture grazed intensively throughout the entire year. In each pasture, we harvested the aboveground biomass from 80 or 100 quadrats of 0.01 m² along a transect and classified the contents by species. We observed 15.5–19.7 species per 0.01 m², which is high richness per 0.01 m² on a worldwide scale. The species richness in the two winter grazing pastures was higher than that in the whole-year grazing pasture. The spatial variation in species richness and species composition in the two winter grazing pastures in which species richness was high was greater than that in the whole-year grazing pasture in which species richness was lower. Most of the leaves that are preserved on the winter grazing pastures during summer are blown away by strong winds during winter, and the remaining leaves are completely exhausted in winter by livestock grazing. A pasture with a high richess is accompanied by a high spatial variation in species richness and species composition. There is a high possibility that the characteristic of spatial variation is also caused by traditional grazing practices in this area.     

Studies on the Transfer of Copper from Soil to Pastures at Different Sampling Periods: A Case Study of a Semiarid Region (Sargodha) in Pakistan

The present investigation was carried out to assess the transfer of copper element from the soil to forage plants consumed by the ruminants in two different pastures at the Livestock Experimental Station at Sargodha, Punjab, Pakistan. Soil and forage samples were collected periodically from two different pastures and analyzed after wet digestion. The survey of copper flow from forage from both pastures in the grazing period exhibited a consistent pattern of decrease from sampling periods 1 to 4 across all the sampling periods. In the legumes and grass pastures, it was decreased regularly and reduced up to 50% to that at the beginning across all the samplings. The copper concentration was higher in the legume pasture than that of grass pasture and sufficient to fulfill the requirement of grazing animals, while in grass pasture, it was higher at the first two sampling periods but dropped to a marginal deficient level at sampling period 3 and reached at the severe deficient level at the fourth sampling period during this investigation. The soil–plant transfer factor for Cu was higher in legume pasture compared to its counterpart. It was found that with the increase of forage maturity, a significant reduction in the forage Cu concentration was observed reaching its minimum level at the last sampling period in the grass pasture. These concentrations were within the marginal and severe deficient levels and provide for only 76% of the ruminant requirements. The naturally upset balance of Cu offers a potential hazard not only to both pastures, but also to the Cu status of grazing ruminants therein. This necessitates the provision of additional amount of Cu mixture in the nutrition of livestock for health and reproduction potential enhancement of the animals being reared at that farm. Supplementing the deficient mineral with locally available Cu feed sources like green fodders, cakes, and brans or providing region-specific mineral supplements would alleviate the deficiency of copper during the late season at the livestock farm.     

Microsatellite analysis of clonality and individual heterozygosity in natural populations of aspen <Emphasis Type="Italic">Populus tremula</Emphasis> L.: Identification of highly heterozygous clone

Aspen Populus tremula L. (Salicaceae) is the fast-growing tree species of environmental and economic value. Aspen is capable of reproduction by both seeds and vegetative means, forming root sprouts. In an adult stand, identification of ramets of one clone among the trees of seed origin based on their morphology is difficult. A panel of 14 microsatellite loci developed for individual identification of aspen was applied for the clonal structure analysis in four natural aspen stands of the European part of Russia: Moscow and Voronezh oblasts, the Mari-El Republic, and the Republic of Tatarstan. In 52 trees from the Moscow sample, 41 multilocus genotypes were identified; in the Voronezh sample, among 30 individuals, 25 different genotypes were detected; and in the sample from Mari-El, 32 trees were represented by 13 genotypes. In the stand from Sabinsky Forestry, Tatarstan, all of the examined 29 trees were represented by a single genotype. The ancestral tree carrier of this genotype which was the most heterozygous (0.929) among all studied aspen individuals (sample mean, 0.598) obviously has spread over a large territory during several cutting and reproduction cycles, currently occupying the area of 2.2 ha. For aspen, usually suffering from Aspen trunk rot, such high viability is evidence of resistance to the main pathogens. The revealed superclone deserves further study with karyological methods and flow cytometry to determine ploidy level and analysis of the growth rate and the quality of wood for possible use in plantation forest production.     

Grazing as a mediator for maintenance of offspring diversity: Sexual and clonal recruitment in alpine grassland communities

To understand the effects of grazing on grassland plants sexual and clonal recruitment, we conducted a demographic field investigation of species recruitment along a grazing gradient in the Tibetan alpine grassland. Grazing intensity had significant effects on quantity and diversity of sexual and clonal recruitment. Sexual recruitment increased significantly, but clonal offspring production decreased significantly with increased grazing intensity. Grazing intensity had different, significant effects on offspring recruitment of the various functional groups in the community, grasses (GG), sedges (SG), legumes (LG) and forbs (FG). Higher grazing intensity reduced offspring recruitment of GG and SG; it increased offspring recruitment of LG and FG. Seedlings were significantly more abundant in lightly grazed, moderately grazed and heavily grazed meadows than in non-grazed grasslands. Offspring diversity from sexual recruitment was significantly higher than that from clonal recruitment in grazed than in non-grazed grasslands. Our studies indicate that moderate grazing had positive effects on seedling recruitment and offspring diversity, but heavy gazing may alter community succession by affecting recruitment patterns among the four plant functional groups.     

Intraspecific competition and light effect on reproduction of Ligularia virgaurea,an invasive native alpine grassland clonal herb

The relationship between sexual reproduction and clonal growth in clonal plants often shows up at the ramet level. However, only a few studies focus on the relationship at the genet level, which could finally account for evolution. The sexual reproduction and clonal growth of Ligularia virgaurea, a perennial herb widely distributed in the alpine grasslands of the Qinghai‐Tibetan Plateau of China, were studied under different competition intensities and light conditions at the genet level through a potted experiment. The results showed that: (1) sexual reproduction did not depend on density or light, and increasing clonal growth with decreasing density and increasing light intensity indicated that intraspecific competition and light intensity may affect the clonal life history of L. virgaurea; (2) both sexual reproduction and clonal growth show a positive linear relationship with genet size under different densities and light conditions; (3) a threshold size is required for sexual reproduction and no evidence of a threshold size for clonal growth under different densities and light conditions; (4) light level affected the allocation of total biomass to clonal and sexual structures, with less allocation to clonal structures and more allocation to sexual structures in full sunlight than in shade; (5) light determined the onset of sexual reproduction, and the genets in the shade required a smaller threshold size for sexual reproduction to occur than the plants in full sunlight; and (6) no evidence was found of trade‐offs between clonal growth and sexual reproduction under different densities and light conditions at the genet level, and the positive correlation between two reproductive modes indicated that these are two integrated processes. Clonal growth in this species may be viewed as a growth strategy that tends to maximize genet fitness.