Carbon source for reproduction in a spring ephemeral herb, Corydalis ambigua (Papaveraceae)

1.  The carbon source for reproduction in plants may differ between flowering and fruiting stages. To clarify how spring ephemerals use current photosynthetic products for reproduction, the allocation patterns of photosynthate at flowering and fruiting and the effects of resource limitation on reproductive performance in Corydalis ambigua were assessed.
2.  A ¹³C tracing experiment revealed that about 20% of the current photosynthetic carbon was used for reproduction at both flowering and fruiting. The proportion of ¹³C allocated to fruits was constant irrespective of the light level. In contrast, ¹³C translocation to tubers increased at fruiting, and this trend was accelerated when plants were shaded.
3.  Defoliation treatment significantly reduced nectar production and tuber mass, while seed production was not affected. Therefore, when carbon assimilation was limited, carbon was preferentially allocated to current reproduction (seeds) rather than to pollinator attraction (nectar) or storage (tuber).
4.  If seed production is partly supported by carbohydrate reserved in the old tissue of tubers, nectar and seed production may not compete strongly for carbon sources. In contrast to the ability of high seed production, the susceptibility of nectar production to current photosynthesis indicates that seed production of this species is basically limited by pollen capture.
5.  Therefore, temporal separation of resource pool for reproduction may mitigate the joint limitation of seed production between pollinator attraction and resource availability. Temporal variation of the sink–source balance of storage organ is crucial to understand the cost of reproduction in perennial plants.     

新疆短命植物小拟南芥(Arabidopsis pumila)种子萌发特性及其生态适应性

通过显微结构及不同处理条件下种子萌发率的观察,对早春短命植物小拟南芥(Arabidopsis pumila)种子萌发特性及影响因素进行了研究,并对其生态适应性进行了讨论。结果表明:(1)温度和光照变化对自然生境和温室收获种子的萌发率影响均不显著,说明此种群在前期萌发阶段对光、温不敏感;(2)自然生境中采收的小拟南芥种子萌发率显著低于温室收获种子,说明环境条件的变化对短命植物种子的发育具有重要作用,可显著改变种子的萌发行为;(3)赤霉素使自然生境收获种子胚活性增强从而对萌发有较大促进作用,可使萌发率增加50%以上;(4)对种皮进行各种机械损伤处理使得种皮松弛或透气,可以显著提高自然生境种子的萌发率(超过70%);(5)盐和干旱胁迫对种子萌发均具有明显的抑制作用,但复水后部分被抑制种子可重新萌发,显示盐和干旱胁迫可导致种子产生浅度休眠。结合小拟南芥自然生存环境及本研究的结果,显示其种子萌发特性与生境具有高度适应性。     

Linking plant spatial aggregation with reproductive traits and near-source seed dispersal: ecological adaptation to heavy grazing

植物的生态适应策略一直备受生态学者的关注,然而很少有研究能够整体地、系统地揭示植物适应草食动物的生态过程。本研究以内蒙古荒漠草原建群种短花针茅为研究对象,分析了重度放牧和不放牧处理下短花针茅的繁殖个体性状、种子性状以及土壤种子库和种群空间格局。研究结果表明,生殖个体性状方面,生殖枝数量对营养枝数量有正向影响;土壤种子库方面,短花针茅生殖个体基部的土壤种子库中短花针茅密度显著高于旁边裸地土壤种子库中短花针茅的密度,且与种子性状和裸地土壤种子库显著负相关;在种群空间格局方面,重度放牧下短花针茅种群空间聚集分布。我们的结果认为在重度放牧处理下,生殖活动在短花针茅的资源分配中处于主导地位;短花针茅植物种群通过调控种子形态特征,采用就近扩散的生态策略,促进了幼年种群与成年种群在较小的尺度上空间正关联,从而形成“安全岛”。     

The evolution of an annual life cycle in killifish: adaptation to ephemeral aquatic environments through embryonic diapause

An annual life cycle is characterized by growth, maturity, and reproduction condensed into a single, short season favourable to development, with production of embryos (seeds, cysts, or eggs) capable of surviving harsh conditions which juveniles or adults cannot tolerate. More typically associated with plants in desert environments, or temperate‐zone insects exposed to freezing winters, the evolution of an annual life cycle in vertebrates is fairly novel. Killifish, small sexually dimorphic fishes in the Order Cyprinodontiformes, have adapted to seasonally ephemeral water bodies across much of Africa and South America through the independent evolution of an annual life history. These annual killifish produce hardy desiccation‐resistant eggs that undergo diapause (developmental arrest) and remain buried in the soil for long periods when fish have perished due to the drying of their habitat. Killifish are found in aquatic habitats that span a continuum from permanent and stable to seasonal and variable, thus providing a useful system in which to piece together the evolutionary history of this life cycle using natural comparative variation. I first review adaptations for life in ephemeral aquatic environments in killifish, with particular emphasis on the evolution of embryonic diapause. I then bring together available evidence from a variety of approaches and provide a scenario for how this annual life cycle evolved. There are a number of features within Aplocheiloidei killifish including their inhabitation of marginal or edge aquatic habitat, their small size and rapid attainment of maturity, and egg properties that make them particularly well suited to the colonization of ephemeral waters.     

Cost of reproduction in a spring ephemeral species, Adonis ramosa (Ranunculaceae): carbon budget for seed production

BACKGROUND AND AIMS: Spring ephemerals have a specific life-history trait, i.e. shoot growth and sexual reproduction occur simultaneously during a short period from snowmelt to canopy closure in deciduous forests. The aim of this study is to clarify how spring ephemerals invest resources for seed production within a restricted period. METHODS: In order to evaluate the cost of reproduction of a typical spring ephemeral species, Adonis ramosa, an experiment was conducted comprising defoliation treatments (intact, one-third and two-thirds leaf-cutting) and fruit manipulations (control, shading and removal) over two growing seasons. In addition, measurements were made of the movements of carbon assimilated via (13)C tracing. KEY RESULTS: Survival rate was high irrespective of treatments and manipulations. The proportion of flowering plants and plant size decreased as a result of the defoliation treatments over 2 years, but the fruit manipulations did not affect flowering activity or plant size. Seed set and seed number decreased as a result of fruit shading treatment, but the defoliation treatments did not affect current seed production. Individual seed weight also decreased in the second year due to fruit shading. The (13)C tracing experiment revealed that young fruits had photosynthetic ability and current photosynthetic products from leaves were mainly transferred to the below-ground parts, while translocation to fruit was very small even when fruit photosynthesis was restricted by the shading treatment. CONCLUSIONS: Current foliage photosynthetic products are largely stored in the below-ground parts for survival and future growth, and about one-third of the resources for seed production may be attained by fruit photosynthesis. Therefore, the trade-off between current seed production and subsequent growth is weak. The cost of seed production may be buffered by sufficient storage in the below-ground organs, effective photosynthesis under high irradiation and self-assimilation ability of fruits.     

克隆植物芦苇叶片功能性状对异质环境的响应

芦苇叶片功能性状的空间变化反映克隆植物的资源分配格局,而其与土壤环境因子的耦合关系体现了克隆植物对异质环境的生态适应策略。本研究以中国西北内陆湿地克隆植物芦苇为对象,分析了湿生生境、盐沼生境、荒漠生境条件下芦苇叶片功能性状及其对土壤环境因子的响应。结果表明: 从湿生生境到荒漠生境,芦苇叶片C、N、P含量分别下降7.2%、40.0%、64.1%,N、P利用效率增加,芦苇叶长、叶宽、叶面积、叶干重、比叶面积和叶厚度均表现出减小趋势。芦苇叶片功能性状间存在协同变化的特征,比叶面积与叶片营养元素表现出显著相关关系。土壤容重、盐分和水分分别是驱动湿生生境、盐沼生境和荒漠生境芦苇叶片功能性状变异的最重要的环境因子。     

延河流域森林草原区不同植物功能型适应策略及功能型物种数量随退耕年限的变化

不同退耕年限退耕地的环境差异以及不同生物间的相互作用导致各阶段植物功能型物种数量不同。研究退耕地植被自然恢复过程中不同植物功能型适应策略及功能型物种数量随退耕年限的变化,对于理解植物对环境的响应机制及植物的适应策略具有重要意义。采用空间序列代替时间序列的方法,以延河流域森林草原区不同退耕年限、自然恢复的植物群落为研究对象,调查了不同退耕年限的植物群落33个,共44种植物,涉及16个科35个属,分别测定了每个物种的叶厚度、比叶面积、叶组织密度、叶片氮含量、比根长、根组织密度、细根氮含量等7项能够反映植物生存对策且易于测量的功能性状。依据这7项植物功能性状,采用数量分类方法将全部物种划分为3个功能型。结果表明:(1)根据C-S-R理论,功能型Ⅰ植物用于防御的投资较多,生长速率处于中间水平,偏向于"胁迫-干扰型",功能型Ⅱ植物能够通过维持体内的养分平衡的方式对抗资源贫瘠或干旱的环境,偏向于"胁迫-竞争型"对策,而功能型Ⅲ植物吸收大量的营养和资源用于生长,偏向于"竞争型";(2)功能型Ⅰ在整个植被恢复时间序列中占据优势地位(61%—80%),并呈增加趋势,功能型Ⅱ则由恢复初期的25%降低为恢复后期的15%,功能型Ⅲ从恢复初期的14%降低到恢复后期的5%。同时,在功能型Ⅰ内部,优势物种也发生着相应的相互替代。虽然土壤养分含量整体上随着植被自然恢复时间的延长而呈上升趋势,但是植物的生存环境并未改善到不存在干扰与胁迫的程度。因此,在植被恢复初期的四、五十年内,"胁迫-干扰型"策略的植物占据着绝对优势。随着植被恢复时间的延长,能够高效利用资源且抗胁迫能力强的物种代替了以快速生长和传播为适应策略的物种。     

Light limitation and litter of an invasive clonal plant,Wedelia trilobata,inhibit its seedling recruitment

Background and Aims

Invasive clonal plants have two reproduction patterns, namely sexual and vegetative propagation. However, seedling recruitment of invasive clonal plants can decline as the invasion process proceeds. For example, although the invasive clonal Wedelia trilobata (Asteraceae) produces numerous seeds, few seedlings emerge under its dense population canopy in the field. In this study it is hypothesized that light limitation and the presence of a thick layer of its own litter may be the primary factors causing the failure of seedling recruitment for this invasive weed in the field.

Methods

A field survey was conducted to determine the allocation of resources to sexual reproduction and seedling recruitment in W. trilobata. Seed germination was also determined in the field. Effects of light and W. trilobata leaf extracts on seed germination and seedling growth were tested in the laboratory.

Key Results

Wedelia trilobata blooms profusely and produces copious viable seeds in the field. However, seedlings of W. trilobata were not detected under mother ramets and few emerged seedlings were found in the bare ground near to populations. In laboratory experiments, low light significantly inhibited seed germination. Leaf extracts also decreased seed germination and inhibited seedling growth, and significant interactions were found between low light and leaf extracts on seed germination. However, seeds were found to germinate in an invaded field after removal of the W. trilobata plant canopy.

Conclusions

The results indicate that lack of light and the presence of its own litter might be two major factors responsible for the low numbers of W. trilobata seedlings found in the field. New populations will establish from seeds once the limiting factors are eliminated, and seeds can be the agents of long-distance dispersal; therefore, prevention of seed production remains an important component in controlling the spread of this invasive clonal plant.     

Rapid and repeatable host plant shifts drive reproductive isolation following a recent human-mediated introduction of the apple maggot fly,Rhagoletis pomonella

Ecological speciation via host-shifting is often invoked as a mechanism for insect diversification, but the relative importance of this process is poorly understood. The shift of Rhagoletis pomonella in the 1850s from the native downy hawthorn, Crataegus mollis, to introduced apple, Malus pumila, is a classic example of sympatric host race formation, a hypothesized early stage of ecological speciation. The accidental human-mediated introduction of R. pomonella into the Pacific Northwest (PNW) in the late 1970s allows us to investigate how novel ecological opportunities may trigger divergent adaptation and host race formation on a rapid timescale. Since the introduction, the fly has spread in the PNW, where in addition to apple, it now infests native black hawthorn, Crataegus douglasii, and introduced ornamental hawthorn, Crataegus monogyna. We use this “natural experiment” to test for genetic differentiation among apple, black, and ornamental hawthorn flies co-occurring at three sympatric sites. We report evidence that populations of all three host-associations are genetically differentiated at the local level, indicating that partial reproductive isolation has evolved in this novel habitat. Our results suggest that conditions suitable for initiating host-associated divergence may be common in nature, allowing for the rapid evolution of new host races when ecological opportunity arises.     

Evolutionary genetics of the aphid Cryptomyzus, with a preliminary analysis of the inheritance of host plant preference, reproductive performance and host-alteration

The aphid species Cryptomyzus galeopsidis (Kaltenbach) includes several distinct forms which have different host plant relationships and life cycles. Cross breeding was used to elucidate the taxonomic status of these forms and to investigate the inheritance of host preference, reproductive performance and host-alternation. One of the forms appeared to be a distinct species because of the reduced fitness of the hybrids. Other host-alternating and non host-alternating forms are considered conspecific and represent two life cycle strategies. Reproductive performance is probably controlled polygenically, since hybrids show an intermediate performance. Host preference in hybrids showed some degree of dominance and seemed to be determined by only a few genes. Host-alternation is presumed to be inherited monofactorially. The implications for speciation are discussed.

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Résumé C. galeopsidis Kaltenbach contient plusieurs formes qui ont différentes relations avec des plantes hôtes et des cycles distincts. Des croisements ont permis d'élucider la taxonomie de ces formes et d'étudier l'hérédité de préférences d'hôtes, des performances reproductives et de l'alternance d'hôtes. Une des formes apparaît comme une espèce distincte par suite de la valeur adaptative réduite des hybrides. Les autres formes avec alternance ou non des hôtes sont considérées comme conspécifiques et représentant deux stratégies vitales différentes. Les performances reproductives sont probablement polygéniques puisque les hybrides ont des performances intermédiaires. Les préférences d'hôtes des hybrides montrent certains degrés de dominance et semblent déterminées par quelques gènes seulement. L'alternance des hôtes est envisagée comme ayant une hérédité monofactorielle. Les conséquences sur la spéciation sont discutées.

     

Oviposition strategy as a means of local adaptation to plant defence in native and invasive populations of the viburnum leaf beetle

Herbivores have been hypothesized to adapt locally to variation in plant defences and such adaptation could facilitate novel associations in the context of biological invasions. Here, we show that in the native range of the viburnum leaf beetle (VLB, Pyrrhalta viburni), two populations of geographically isolated hosts-Viburnum opulus and Viburnum tinus-have divergent defences against VLB oviposition: negative versus positive density-dependent egg-crushing wound responses, respectively. Populations of beetles coexisting with each host show an adaptive behavioural response: aggregative versus non-aggregative oviposition on V. opulus and V. tinus, respectively. In parallel, we show that in North America, where VLB is invasive, defences of three novel hosts are negatively density-dependent, and beetles' oviposition behaviour is aggregative. Thus, local adaptation to plant defences has the potential to facilitate the invasion of herbivores onto novel hosts.     

Conservation genetics and breeding system of Penstemon debilis (Plantaginaceae), a rare beardtongue endemic to oil shale talus in western Colorado,USA

Rare species of plants are especially vulnerable to extinction when populations are few, have small numbers of individuals, and are fragmented. Such conditions lead to a reduction in gene flow and genetic diversity, and encourage inbreeding depression. We conducted a study of the reproductive biology and population genetics of Penstemon debilis (Plantaginaceae), a Federally Threatened species endemic to a small region of oil shale extraction in western Colorado, USA. Most of the habitat area is privately owned and undergoing natural gas extraction activities. Penstemon debilis reproduces both vegetatively and as an outcrosser that requires a pollen vector. Moderate levels of inbreeding, but no inbreeding depression, were found within populations of P. debilis. Genetic divergence among the extant populations surveyed was moderate (F_ST values = 0.069–0.231; Nm = 0.831–3.385) with levels of genetic diversity within populations relatively low compared to congeners with similar modes of pollination and reproductive biology. STRUCTURE analysis revealed three population clusters with some admixture among all extant populations. Genetic diversity within and among P. debilis populations is similar to genetic diversity found for other rare and endemic outcrossing plant species. Our results are consistent with a pattern of recent population fragmentation or low levels of pollen‐mediated gene flow among populations in close proximity to one another. Conservation of P. debilis will require cooperative management strategies between private landowners, government agencies, and concerned NGOs to preserve habitat for this rare species.     

Temporal and host-related variation in frequencies of genes that enable Phyllotreta nemorum to utilize a novel host plant, Barbarea vulgaris

The flea beetle, Phyllotreta nemorum L. (Coleoptera: Chrysomelidae), is an intermediate specialist feeding on a small number of plants within the family Brassicaceae. The most commonly used host plant is Sinapis arvensis L., whereas the species is found more rarely on Cardaria draba (L.) Desv., Barbarea vulgaris R.Br., and cultivated radish (Raphanus sativus L.). The interaction between flea beetles and Barbarea vulgaris ssp. arcuata (Opiz.) Simkovics seems to offer a good opportunity for experimental studies of coevolution. The plant is polymorphic, as it contains one type (the P‐type) that is susceptible to all flea beetle genotypes, and another type (the G‐type) that is resistant to some genotypes. At the same time, the flea beetle is also polymorphic, as some genotypes can utilize the G‐type whereas others cannot. The ability to utilize the G‐type of B. vulgaris ssp. arcuata is controlled by major dominant genes (R‐genes). The present investigation measured the frequencies of flea beetles with R‐genes in populations living on different host plants in 2 years (1999 and 2003). Frequencies of beetles with R‐genes were high in populations living on the G‐type of B. vulgaris ssp. arcuata in both years. Frequencies of beetles with R‐genes were lower in populations living on other host plants, and declining frequencies were observed in five out of six populations living on S. arvensis. Selection in favour of R‐genes in populations living on B. vulgaris is the most likely mechanism to account for the observed differences in the relative abundance of R‐genes in flea beetle populations utilizing different host plants. A geographic mosaic with differential levels of interactions between flea beetles and their host plants was demonstrated.     

Recursive adaptation in action: allochronic isolation and divergence of host-associated populations of the apple maggot fly,Rhagoletis pomonella,following its recent introduction to the western USA

An outstanding issue in the study of insect host races concerns the idea of ‘recursive adaptive divergence’, whereby adaptation can occur repeatedly across space and/or time, and the most recent adaptive episode is defined by one or more previously similar cases. The host plant shift of the apple maggot fly, Rhagoletis pomonella (Walsh) (Diptera: Tephritidae, Carpomyini), from ancestral downy hawthorn [Crataegus mollis (Torr. & A. Gray) Scheele] to introduced, domesticated apple (Malus domestica Borkh.) in the eastern USA has long served as a model system for investigating ecologically driven host race formation in phytophagous insect specialists. Here, we report results from an annual geography survey of eclosion time demonstrating a similar ecological pattern among nascent host-associated populations of the fly recently introduced ca. 40 years ago from its native range in the east into the Pacific Northwest (PNW) region of the USA. Specifically, using data collected from 25 locations across 5 years, we show that apple-infesting fly populations in the PNW have rapidly and repeatedly shifted (and maintained differences in) their adult eclosion life-history timing to infest two novel hawthorn hosts with different fruiting phenologies – a native species (Crataegus douglasii Lindl.) and an introduced species (Crataegus monogyna Jacq.) – generating partial allochronic reproductive isolation in the process. The shifts in the PNW parallel the classic case of host race formation in the eastern USA, but have occurred bi-directionally to two hawthorn species with phenologies slightly earlier (black hawthorn) and significantly later (ornamental hawthorn) than apple. Our results imply that R. pomonella can both possess and retain extensive-standing variation (i.e., ‘adaptive memory’) in diapause traits, even following introductions, to rapidly and temporally track novel phenological host opportunities when they arise. Thus, ‘specialized’ host races may not constitute evolutionary dead ends. Rather, adaptive phenotypic and genetic memory may carry over from one host shift to the next, recursively facilitating host race formation in phytophagous insects.