Frequency of cyanogenesis in tropical rainforests of far north Queensland, Australia

BACKGROUND AND AIMS: Plant cyanogenesis is the release of toxic cyanide from endogenous cyanide-containing compounds, typically cyanogenic glycosides. Despite a large body of phytochemical, taxonomic and ecological work on cyanogenic species, little is known of their frequency in natural plant communities. This study aimed to investigate the frequency of cyanogenesis in Australian tropical rainforests. Secondary aims were to quantify the cyanogenic glycoside content of tissues, to investigate intra-plant and intra-population variation in cyanogenic glycoside concentration and to appraise the potential chemotaxonomic significance of any findings in relation to the distribution of cyanogenesis in related taxa. METHODS: All species in six 200 m(2) plots at each of five sites across lowland, upland and highland tropical rainforest were screened for cyanogenesis using Feigl-Anger indicator papers. The concentrations of cyanogenic glycosides were accurately determined for all cyanogenic individuals. KEY RESULTS: Over 400 species from 87 plant families were screened. Overall, 18 species (4.5 %) were cyanogenic, accounting for 7.3 % of total stem basal area. Cyanogenesis has not previously been reported for 17 of the 18 species, 13 of which are endemic to Australia. Several species belong to plant families or orders in which cyanogenesis has been little reported, if at all (e.g. Elaeocarpaceae, Myrsinaceae, Araliaceae and Lamiaceae). A number of species contained concentrations of cyanogenic glycosides among the highest ever reported for mature leaves-up to 5.2 mg CN g(-1) d. wt, for example, in leaves of Elaeocarpus sericopetalus. There was significant variation in cyanogenic glycoside concentration within individuals; young leaves and reproductive tissues typically had higher cyanogen content. In addition, there was substantial variation in cyanogenic glycoside content within populations of single species. CONCLUSIONS: This study expands the limited knowledge of the frequency of cyanogenesis in natural plant communities, includes novel reports of cyanogenesis among a range of taxa and characterizes patterns in intra-plant and intra-population variation of cyanogensis.     

Spectrophotometer-aided evaluation of cyanogenic potential in white clover (Trifolium repens L.)

In comparison with ordinary methods of colorimetric evaluation of cyanogenic potential based on visual evaluation of the alkaline picrate reaction, a spectrophotometer-aided method could be more accurate (since it determines the exact amount of hydrogen cyanide released by the plant material), and less time-consuming as it can be performed on bulk material rather than on a number of individual plants. Ten white clover populations were evaluated by a spectrophotometer-aided method and by two visual evaluation criteria. All methods indicated the presence of large variation between populations. Visual methods gave almost identical results and allowed only for the distinction between cyanogenic and substantially acyanogenic populations. The results were only moderately consistent with those obtained by the spectrophotometer-aided method, which could detect the presence of variation also between cyanogenic populations. The effects of various incubation times (from 4 to 48 h) and of the addition of β-glucosidase on hydrogen cyanide release were also investigated. Comparable results for ranking of populations could be obtained over a range of incubation times, but at least 24 h were needed for a reliable estimation of the hydrogen cyanide produced by plants. The addition of enzyme did not increase the released cyanide. The effect of season and/or conditions of evaluation was marked on mean cyanogenic potential but limited on ranking of populations. Copyright © 2000 John Wiley & Sons, Ltd.     

Variation of cyanogenesis in some plant species of the midwestern United States

The frequency of cyanogenesis of 48 species of vascular plants was examined by testing 30 individuals from five populations of each species for release of cyanide. The rate at which cyanide was released and the amount of cyanide released varied widely among individuals of a population and among populations of a species. For many taxa, the frequency of cyanogenesis was highly variable among populations. Of the species examined, 20 have not been reported previously as being cyanogenic.     

Positive effects of cyanogenic glycosides in food plants on larval development of the common blue butterfly

Cyanogenesis is a widespread chemical defence mechanism in plants against herbivory. However, some specialised herbivores overcome this protection by different behavioural or metabolic mechanisms. In the present study, we investigated the effect of presence or absence of cyanogenic glycosides in birdsfoot trefoil (Lotus corniculatus, Fabaceae) on oviposition behaviour, larval preference, larval development, adult weight and nectar preference of the common blue butterfly (Polyommatus icarus, Lycaenidae). For oviposition behaviour there was a female-specific reaction to cyanogenic glycoside content; i.e. some females preferred to oviposit on cyanogenic over acyanogenic plants, while other females behaved in the opposite way. Freshly hatched larvae did not discriminate between the two plant morphs. Since the two plant morphs differed not only in their content of cyanogenic glycoside, but also in N and water content, we expected these differences to affect larval growth. Contrary to our expectations, larvae feeding on cyanogenic plants showed a faster development and stronger weight gain than larvae feeding on acyanogenic plants. Furthermore, female genotype affected development time, larval and pupal weight of the common blue butterfly. However, most effects detected in the larval phase disappeared for adult weight, indicating compensatory feeding of larvae. Adult butterflies reared on the two cyanogenic glycoside plant morphs did not differ in their nectar preference. But a gender-specific effect was found, where females preferred amino acid-rich nectar while males did not discriminate between the two nectar mimics. The presented results indicate that larvae of the common blue butterfly can metabolise the surplus of N in cyanogenic plants for growth. Additionally, the female-specific behaviour to oviposit preferably on cyanogenic or acyanogenic plant morphs and the female-genotype-specific responses in life history traits indicate the genetic flexibility of this butterfly species and its potential for local adaptation.     

Detection of selection in populations of white clover (Trifolium repens L.)

Seed populations of white clover polymorphic for the presence/absence of both ovariogenic glucosides and the hydrolysing enzyme linamarase, were introduced into three natural populations. Over the first six months of life a significant increase in the frequency of linamarase containing individuals occurred. Estimated selection coefficients against plants lacking linamarase were in the region of 0.3. This result may have been due to selection at the enzyme locus alone, or to selection favouring cyanogenic individuals which possess both cyanogenic glucosides and enzyme.     

Influence of two selective factors on cyanogenesis polymorphism ofTrifolium repens L. in Darjeeling Himalaya

Cyanogenesis-the production of toxic hydrogen cyanide (HCN) by damaged tissue-inTrifolium repens L. (white clover), a type of most important pasture legume, has been studied at different elevations of Darjeeling Himalaya (latitude-27° 2′ 57″ N, longitude-88° 15′ 45″ E). Release of HCN takes place due to reaction between cyanogenic glucosides stored in vacuoles of the leaf cell and the corresponding enzyme β-glucosidase present in another compartment, often cell wall. Cyanogenesis, a defense system in plant, protects the clover from herbivore and inhibits grazing. Biochemical analysis showed the presence and absence of the cyanogenesis trait within the population in different proportions at different elevations. Acyanogenic individuals also showed variations with respect to presence or absence of either cyanogenic glucosides or β-glucosidase enzyme or both. The distribution of cyanogenic and acyanogenic plants was found in all places, but at lower altitudes (2084–2094 m) the dominating plants were cyanogenic whereas in higher altitude (2560 m) the dominating plants were acyanogenic. It was observed that blister beetle (Mylabris pustalata Thunb.) and the mollusc (Macrochlamys tusgurium Benson.) were the most common consumer of leaflets ofT. repens. Six categories of damage on white clover leaf by these animals were recorded. Our results suggest that the two selective factors or forces i.e. very cold temperature (harmful to cyanogenic plants) at higher altitude as well as indiscriminate but preferential predation (harmful to acyanogenic plants) interact to affect the system of cyanogenesis and also to cause the stable and protective polymorphism inT. repens rather than genotypic differences present among the plants.     

The lack of genetic bottleneck in invasive Tansy ragwort populations suggests multiple source populations.

Jacobaea vulgaris (Asteraceae) is a species of Eurasian origin that has become a serious non-indigenous weed in Australia, New Zealand, and North America. We used neutral molecular markers to (1) test for genetic bottlenecks in invasive populations and (2) to investigate the invasion pathways. It is for the first time that molecular markers were used to unravel the process of introduction in this species.The genetic variation of 15 native populations from Europe and 16 invasive populations from Australia, New Zealand and North America were compared using the amplified fragment length polymorphisms (AFLP's). An analysis of molecular variance showed that a significant part (10%) of the total genetic variations between all individuals could be explained by native or invasive origin.Significant among-population differentiation was detected only in the native range, whereas populations from the invasive areas did not significantly differ from each other; nor did the Australian, New Zealand and North American regions differ within the invasive range. The result that native populations differed significantly from each other and that the amount of genetic variation, measured as the number of polymorphic bands, did not differ between the native and invasive area, strongly suggests that introductions from multiple source populations have occurred. The lack of differentiation between invasive regions suggests that either introductions may have occurred from the same native sources in all invasive regions or subsequent introductions took place from one into another invasive region and the same mix of genotypes was subsequently introduced into all invasive regions.An assignment test showed that European populations from Ireland, the Netherlands and the United Kingdom most resembled the invasive populations.     

Genetic diversity and structure of wild populations of the tropical dry forest tree Jacaratia mexicana (Brassicales: Caricaceae) at a local scale in Mexico

The tropical dry forest is a greatly endangered ecosystem, from which Jacaratia mexicana is a native tree. With the aim to assess the levels of genetic variation and population structure, four wild populations of J. mexicana were studied in the Sierra de Huautla Biosphere Reserve, Morelos, Mexico. For this, DNA was extracted from 159 individuals and were amplified with six random primers using the Random Amplified Polymorphic DNA (RAPD). A total of 54 bands were obtained, of which 50 (92.6%) were polymorphic. The total genetic diversity found within the four populations was 0.451 when estimated by Shannon's index. An AMOVA analysis showed that 84% of the total genetic variation was found within populations and 16% was among populations. The UPGMA dendrogram showed that all individuals from one of the populations (Huaxtla) formed one distinct genetic group, while the rest of the individuals did not cluster according to population. A Mantel test did not show an association between genetic and geographical distances among populations (r=0.893, p=0.20). A Bayesian cluster analysis performed with STRUCTURE, showed that the most probable number of genetic groups in the data was four (K=4), and confirmed the distinctness of Huaxtla population. Our results showed that important genetic differentiation among populations can occur even at this small geographic scale and this has to be considered in conservation actions for this genetic resource.     

Cyanogenic constituents in woody plants in natural lowland rain forest in Costa Rica

Screening of fresh leaves and other plant organs revealed the presence of cyanogenesis in 25 species in 16 families out of a total of 488 species in 79 families vouchered in natural lowland rain forest in Costa Rica. In a qualitative screening of a random sample based on seven one-hectare inventories of woody plants, we found cyanogenesis in 4.0% of all species; these represented 2.5% of all individuals present, and 3.0% of the total basal area of stems. The frequency of occurrence of cyanogenic compounds was higher in reproductive plant parts than in leaves, and highest in pericarps. Cyanogenesis was found in Annona amazonica, Annona pittieri, Cymbopetalum costaricense, Tabebuia chrysantha, Sloanea tuerckheimii, Sapium laurifolium, Lecointea amazonica, Carpotroche platyptera, Klayana odorata, Byrsonima crispa, Miconia splendens, Inga acuminata, Chaunochiton kappleri, Passiflora ambigua, P. pittieri, P. talamancensis, P. vitifolia, Panopsis costaricensis, Faramea pawibractea, Paullinia capreolata, Pouteria amygdalicarpa, P. campechiana, P. subrotata, P. torta and Rinorea guatemalensis. Some taxonomic implications are discussed. The identity of cyanogenic constituents found in dried material of these species is presented, as is the total cyanogenic potential (CNp) of 11 positive species. The CNp ranged from less than 5 to approx. 2000 mg kg^-1 f.w. Several species were cyanogenic only in some individuals. A total of 463 species gave no positive test for cyanogenic glycosides. Tests on leaves and seeds of Ampelocera macrocarpa indicated the content of a rare and unknown volatile constituent.     

Efficacy of population structure analysis with breeding populations and inbred lines

The objective was to assess by simulation the efficacy of population structure analysis in plant breeding. Twelve populations and 300 inbred lines were simulated and genotyped using 100 microsatellite loci. The experimental material included populations with and without admixture, ancestry relationship and linkage disequilibrium, and with distinct levels of genetic differentiation and effective sizes. The analyses were performed using Structure software and employed all available models. For all the group number (K) tested, for both populations and inbred lines, the admixture model with correlated allelic frequencies provided the highest value for the logarithm of the marginal likelihood. Fitting appropriate model and using adequate sample size for individuals and markers, Structure was effective in identifying the correct population structure, migrants and individuals with genome from distinct populations. The linkage model did not result in an improvement in clustering relative to the admixture model with correlated allelic frequencies. The inclusion of prior information did not change the results; for some K values the analyses showed slight higher values of the marginal likelihood. The reduction in the number of individuals and markers negatively affected the results. There was a high variation in the most probable K value between the evaluated methods.     

不同地区甜菜夜蛾种群的遗传多样性分析

甜菜夜蛾Spodoptera exigua (Hübner)是一种重要的农业害虫,曾给我国农牧业生产造成过重大的经济损失。为了阐明甜菜夜蛾的种群动态规律,改善和提高其预测、防治水平,我们应用扩增片段长度多态性(AFLP)技术对我国7个甜菜夜蛾种群的42头雄蛾的遗传多样性进行了研究分析。结果表明,不同种群个体间遗传相似性指数分布在0.143～0.824之间,同一种群内不同个体间的遗传相似性指数分布在0.250～0.786之间,同一种群内个体间的遗传相似性指数有许多小于不同种群个体间的。江西种群的多态性条带比例最低,为80.7%; 山东种群的多态性条带比例最高,达88.6%。总体而言,北方种群内的遗传多样性高于南方种群。聚类分析结果表明,同一种群的个体并不总能聚在一起,即种群间不存在明显的遗传分化。这些结果为明确我国甜菜夜蛾的迁飞规律及各发生危害区的虫源关系提供了进一步的实验依据。     

Rapid evolution of an adaptive cyanogenesis cline in introduced North American white clover (Trifolium repens L.)

White clover is polymorphic for cyanogenesis (HCN production after tissue damage), and this herbivore defence polymorphism has served as a classic model for studying adaptive variation. The cyanogenic phenotype requires two interacting biochemical components; the presence/absence of each component is controlled by a simple Mendelian gene (Ac/ac and Li/li). Climate-associated cyanogenesis clines occur in both native (Eurasian) and introduced populations worldwide, with cyanogenic plants predominating in warmer locations. Moreover, previous studies have suggested that epistatic selection may act within populations to maintain cyanogenic (AcLi) plants and acyanogenic plants that lack both components (acli plants) at the expense of plants possessing a single component (Acli and acLi plants). Here, we examine the roles of selection, gene flow and demography in the evolution of a latitudinal cyanogenesis cline in introduced North American populations. Using 1145 plants sampled across a 1650 km transect, we determine the distribution of cyanogenesis variation across the central United States and investigate whether clinal variation is adaptive or an artefact of population introduction history. We also test for the evidence of epistatic selection. We detect a clear latitudinal cline, with cyanogenesis frequencies increasing from 11% to 86% across the transect. Population structure analysis using nine microsatellite loci indicates that the cline is adaptive and not a by-product of demographic history. However, we find no evidence for epistatic selection within populations. Our results provide strong evidence for rapid adaptive evolution in these introduced populations, and they further suggest that the mechanisms maintaining adaptive variation may vary among populations of a species.     

Cyanogenic glycosides and menisdaurin from Guazuma ulmifolia, Ostrya virgininana, Tiquilia plicata and Tiquilia canescens

The major cyanogenic glycoside of Guazuma ulmifolia (Sterculiaceae) is (2R)-taxiphyllin (>90%), which co-occurs with (2S)-dhurrin. Few individuals of this species, but occasional other members of the family, have been reported to be cyanogenic. To date, cyanogenic compounds have not been characterized from the Sterculiaceae. The cyanogenic glycosides of Ostrya virginiana (Betulaceae) are (2S)-dhurrin and (2R)-taxiphyllin in an approximate 2:1 ratio. This marks the first report of the identification of cyanogenic compounds from the Betulaceae. Based on NMR spectroscopic and TLC data, the major cyanogenic glucoside of Tiquilia plicata is dhurrin, whereas the major cyanide-releasing compound of Tiquilia canescens is the nitrile glucoside, menisdaurin. NMR and TLC data indicate that both compounds are present in each of these species. The spectrum was examined by CI-MS, ¹H and ¹³C NMR, COSY, 1D selective TOCSY, NOESY, and ¹J/^2,3J HETCOR experiments; all carbons and protons are assigned. The probable absolute configuration of (2R)-dhurrin is established by an X-ray crystal structure. The ¹H NMR spectrum of menisdaurin is more complex than might be anticipated, containing a planar conjugated system in which most elements are coupled to several other atoms in the molecule. The coupling of one vinyl proton to the protons on the opposite side of the ring involves a ⁶J- and a ^5/7J-coupling pathway. A biogenetic pathway for the origin of nitrile glucosides is proposed.     

Cyanogenic Eucalyptus nobilis is polymorphic for both prunasin and specific beta-glucosidases

Cyanogenesis (i.e. the evolution of HCN from damaged plant tissue) requires the presence of two biochemical pathways, one controlling synthesis of the cyanogenic glycoside and the other controlling the production of a specific degradative beta-glucosidase. The sole cyanogenic glycoside in Eucalyptus nobilis was identified as prunasin (D-mandelonitrile beta-D-glucoside) using HPLC and GC-MS. Seedlings from three populations of E. nobilis were grown under controlled conditions and 38% were found to be acyanogenic, a proportion far greater than reported for any other cyanogenic eucalypt. A detailed study of the acyanogenic progeny from a single open-pollinated parent found that 23% lacked a cyanogenic beta-glucosidase, 32% lacked prunasin and 9% lacked both. Of the remaining seedlings initially identified as acyanogenics, 27% contained either trace amounts of beta-glucosidase or prunasin, while 9% contained trace amounts of both. Results support the hypothesis that the two components necessary for cyanogenesis are inherited independently. Trace amounts are likely to result from the presence of non-specific beta-glucosidases or the glycosylation of the cyanohydrin intermediate by non-specific UDP glycosyl transferases.     

Spatial repartition and genetic relationship of green and albino individuals in mixed populations of Cephalanthera orchids

Several green orchids of the Neottieae tribe acquire organic carbon both from their mycorrhizal fungi and from photosynthesis. This strategy may represent an intermediate evolutionary step towards mycoheterotrophy of some non‐photosynthetic (albino) orchids. Mixed populations of green and albino individuals possibly represent a transient evolutionary stage offering opportunities to understand the evolution of mycoheterotrophy. In order to understand the emergence of albinos, we investigated patterns of spatial and genetic relationships among green and albino individuals in three mixed populations of Cephalanthera damasonium and one of C. longifolia using spatial repartition and Amplified fragment length polymorphism (AFLP) markers. Two of these populations were monitored over two consecutive flowering seasons. In spatial repartition analyses, albino individuals did not aggregate more than green individuals. Genetic analyses revealed that, in all sampled populations, albino individuals did not represent a unique lineage, and that albinos were often closer related to green individuals than to other albinos from the same population. Genetic and spatial comparison of genets from the 2‐year monitoring revealed that: (i) albinos had lower survival than green individuals; (ii) accordingly, albinos detected in the first year did not correspond to the those sampled in the second year; and (iii) with one possible exception, all examined albinos did not belong to any green genet from the same and/or from the previous year, and vice versa. Our results support a scenario of repeated insurgence of the albino phenotypes within the populations, but unsuccessful transition between the two contrasting phenotypes. Future studies should try to unravel the genetic and ecological basis of the two phenotypes.     

Aridity shapes cyanogenesis cline evolution in white clover (Trifolium repens L.)

Adaptive differentiation between populations is often proposed to be the product of multiple interacting selective pressures, although empirical support for this is scarce. In white clover, populations show adaptive differentiation in frequencies of cyanogenesis, the ability to produce hydrogen cyanide after tissue damage. This polymorphism arises through independently segregating polymorphisms for the presence/absence of two required cyanogenic components, cyanogenic glucosides and their hydrolysing enzyme. White clover populations worldwide have evolved a series of recurrent, climate‐associated clines, with higher frequencies of cyanogenic plants in warmer locations. These clines have traditionally been hypothesized to reflect a fitness trade‐off between chemical defence in herbivore‐rich areas (warmer climates) and energetic costs of producing cyanogenic components in areas of low herbivore pressure (cooler climates). Recent observational studies suggest that cyanogenic components may also be beneficial in water‐stressed environments. We investigated fitness trade‐offs associated with temperature‐induced water stress in the cyanogenesis system using manipulative experiments in growth chambers and population surveys across a longitudinal precipitation gradient in the central United States. We find that plants producing cyanogenic glucosides have higher relative fitness in treatments simulating a moderate, persistent drought stress. In water‐neutral treatments, there are energetic costs to producing cyanogenic components, but only in treatments with nutrient stress. These fitness trade‐offs are consistent with cyanogenesis frequencies in natural populations, where we find clinal variation in the proportion of plants producing cyanogenic glucosides along the precipitation gradient. These results suggest that multiple selective pressures interact to maintain this adaptive polymorphism and that modelling adaptation will require knowledge of environment‐specific fitness effects.     

Studies on Variability in White Clover: Growth Habits and Cyanogenic Glucosides

Growth habits and cyanogenesis were studied in a field experimentwith white clover (Trifolium repens L.) cv. Huia. Eighty controlplants propagated by seeds, and 80 clones of an in vivo selectedvariant were examined in mid-late August and late September.The temperature effect on cyanogenic glucoside levels was alsoexamined on in vitro grown plantlets of the variant in growthchambers. Results showed that the upright growth habit dominated in controlplants. The population of the variant plants was mainly composedof prostrate clones and a transition from prostrate to uprightgrowth habit occurred. Considerable variation was observed withregard to all measured morphological characters in both controland variant plants. Great variation was also noted in cyanogenicglucoside content in the leaf laminae of the control plants.Low cyanogenic plants, on the other hand, dominated in the variant.The number of low and high cyanogenic plants increased in thecontrol by the end of the growth season, after the first frosts.A one-way shift of cyanogenic (slight increase) was recordedin the variant. Similar levels of cyanogenic glucoside and linamarasewere determined in the in vitro and field-grown plants. Cyanogenicglucoside content slightly decreased with the age of the invitro plantlets, but variation in temperature did not causeany changes in their level. The response to the environmentalchanges, regarding cyanogenesis, appear to be genetically determined. Trifolium repens, in vitro, somaclonal variation, cyanogenic glucosides     

Cyanogenic polymorphism in Eucalyptus polyanthemos Schauer subsp. vestita L. Johnson and K. Hill (Myrtaceae)

Plant cyanogenesis, the release of cyanide from endogenous cyanide-containing compounds, is an effective herbivore deterrent. This paper characterises cyanogenesis in the Australian tree Eucalyptus polyanthemos Schauer subsp. vestita L. Johnson and K. Hill for the first time. The cyanogenic glucoside prunasin ((R)-mandelonitrile β-D-glucoside) was determined to be the only cyanogenic compound in E. polyanthemos foliage. Two natural populations of E. polyanthemos showed quantitative variation in foliar prunasin concentration, varying from zero (i.e. acyanogenic) to 2.07 mg CN g^-1 dry weight in one population and from 0.17 to 1.98 mg CN g^-1 dry weight in the other. No significant difference was detected between the populations with respect to the mean prunasin concentration or the degree of variation in foliar prunasin, despite significant differences in foliar nitrogen. Variation between individuals was also observed with respect to the capacity of foliage to catabolise prunasin to form cyanide. Moreover, variation in this capacity generally correlated with the amount of prunasin in the tissue, suggesting genetic linkage between prunasin and β-glucosidase.     

心叶凹唇姜的开花格局和雄性先熟机制

心叶凹唇姜 (Boesenbergia longiflora) 是分布于亚洲热带的一种姜科植物, 在我国滇南地区成片生长于石灰山贫瘠的岩石缝隙和路边林下的冲积土壤上。心叶凹唇姜的花寿命为两天, 第1天为严格的雄性阶段, 第2天为严格的雌性阶段, 是典型的花内雄性先熟, 一朵花的雌、雄性阶段没有重叠, 因而完全避免了自花授粉的发生。对这两种生境 (石灰山贫瘠的岩石缝隙和路边林下) 下的心叶凹唇姜的4个野外种群 (在云南省普洱市的翠云乡彩虹洞、南岛河、澜沧县惠民乡和西双版纳州绿石林森林公园) 和栽培植株的开花格局进行调查后发现, 大多数个体在同一时间内只有一个花序开花, 并且每隔一天才开一朵花, 只有10%左右的植株同时具有雄性阶段和雌性阶段的花, 这样的开花格局有效地降低了同株异花授粉发生的可能。心叶凹唇姜的平均花粉-胚珠比 (Pollen-ovuleratio) 为291, 应该是以异交为主的混合交配系统。其自然种群存在着花粉限制, 彩虹洞、南岛河和绿石林3个不同种群的平均结籽率都不到60%, 这可能与其较小的花展示有关。心叶凹唇姜为泛化传粉植物, 不同种群的有效传粉昆虫种类各不相同。在两种不同的生境条件下, 心叶凹唇姜花的大小和开花格局都没有显著的差异, 大多数个体同时只具有一种性阶段的花, 说明心叶凹唇姜这种独特的开花格局是长期进化的结果。心叶凹唇姜通过隔一天开一朵花的开花格局, 配合雄性先熟机制, 有效地降低了自交的发生。     

The effects of altitude,aspect, grazing and time on the proportion of cyanogenics in neighbouring populations of Trifolium repens L. (white clover)

The proportion of cyanogenic individuals of white clover amongst 200 individuals in each of 32 neighbouring populations in Northumberland, UK were recorded. Sites differed for four altitude zones, two aspects, and were either sheep pasture or ungrazed meadow. The proportion of cyanogenics showed a striking reduction with increasing altitude. Below 100 m, north-facing sites contained more cyanogenics than south-facing sites, but aspect did not affect higher sites. Grassland management had no effect on the proportion of cyanogenics. A 24-year study of one population showed highly significant changes in the proportion of cyanogenics over time, suggesting that a significant turnover of individuals occurs. In some cases, different size class cohorts varied for the proportion of cyanogenics within a year, and the same cohort varied between years. We conclude that environment at birth may determine the proportion of cyanogenics for that cohort, so that this proportion persists in that cohort as it matures. Comparisons of the proportion of cyanogenics with mean monthly averages for January minimum temperature, July maximum temperature and August rainfall showed a significant association only with mean January minima 2 1/2 years previously. We hypothesise that the 2 to 3-year cohort may predominate in this population. We suggest that winter cold and summer drought may both select against cyanogenics. Grazing by large herbivores does not favour cyanogenesis, but some invertebrate herbivory may do so. Most selection probably occurs at birth and will be greatest in populations with a high turnover.