Herbivory and ramet performance in the clonal herb Trientalis europaea L.

1 The intensity of leaf damage caused by invertebrate herbivores and grazing by vertebrates and their effects on clonal growth, survival and reproduction were examined in a population of the forest herb Trientalis europaea during 4 years.
2 Levels of herbivory were low and varied between years in the studied population. Larger ramets were exposed to a greater risk of both grazing and leaf defoliation. Ramets that suffered leaf damage in 1 year experienced an increased probability of leaf damage in succeeding years. The probability of suffering herbivore attack was independent among the ramets belonging to the same clonal fragment, suggesting that clonal propagation might operate as a risk-spreading strategy in this species.
3 Leaf damage did not affect any measure of plant performance, probably due to the low amounts of leaf area removed by invertebrate herbivores. In contrast, vertebrate grazing affected all phases of the pseudo-annual life cycle of T. europaea . Grazing prevented flowering and fruiting, increased ramet mortality during summer and decreased tuber production. Furthermore, grazed ramets produced shorter stolons and smaller tubers, which in turn had a lower winter survival and produced smaller ramets in the following growing season. The large impact of grazing was due to the consumption of the whole of the single shoot of ramets of T. europaea . Although regrowth was possible, secondary shoots were significantly smaller and assimilation was delayed.
4 Tubers originating from grazed ramets were placed shallower than tubers from ungrazed ramets. This could be due to an alteration in the growth pattern of stolons of the grazed ramets.     

Effects of Heterogeneous Competitor Distribution and Ramet Aggregation on the Growth and Size Structure of a Clonal Plant

Spatially heterogeneous distribution of interspecific competitors and intraspecific aggregation of offspring ramets may affect the growth and size structure of clonal plant populations, but these have been rarely studied. We conducted a greenhouse experiment in which we grew a population of eight offspring ramets (plants) of the stoloniferous clonal plant Hydrocotyle vulgaris aggregately or segregately in two homogeneous treatments with or without a competing grass Festuca elata and a heterogeneous treatment with a patchy distribution of the grass. In patchy grass treatments, H. vulgaris produced markedly more biomass, ramets and stolons in open patches (without grasses) than in grass patches, but displayed lower size variations as measured by coefficient of variation of biomass, ramets and stolons among the eight plants. In open areas, H. vulgaris produced statistically the same amounts of biomass and even more stolons and showed higher size variations in patchy grass treatments than in open (no grass) treatments. In grass areas, H. vulgaris grew much worse and displayed higher size variations in patchy grass treatments than in full grass treatments. Ramet aggregation decreased the growth of H. vulgaris in open treatments and in both open and grass patches in patchy grass treatments, but had little effect in full grass treatments. Ramet aggregation had little effect on size variations. Therefore, heterogeneous distribution of competitors can affect the growth and size structure of clonal plant populations, and ramet aggregation may decrease population growth when they grow in open environments or heterogeneous environments with a patchy distribution of interspecific competitors.     

Impact of herbivores on nitrogen cycling: contrasting effects of small and large species

Herbivores are reported to slow down as well as enhance nutrient cycling in grasslands. These conflicting results may be explained by differences in herbivore type. In this study we focus on herbivore body size as a factor that causes differences in herbivore effects on N cycling. We used an exclosure set-up in a floodplain grassland grazed by cattle, rabbits and common voles, where we subsequently excluded cattle and rabbits. Exclusion of cattle lead to an increase in vole numbers and a 1.5-fold increase in net annual N mineralization at similar herbivore densities (corrected to metabolic weight). Timing and height of the mineralization peak in spring was the same in all treatments, but mineralization in the vole-grazed treatment showed a peak in autumn, when mineralization had already declined under cattle grazing. This mineralization peak in autumn coincides with a peak in vole density and high levels of N input through vole faeces at a fine-scale distribution, whereas under cattle grazing only a few patches receive all N and most experience net nutrient removal. The other parameters that we measured, which include potential N mineralization rates measured under standardized laboratory conditions and soil parameters, plant biomass and plant nutrient content measured in the field, were the same for all three grazing treatments and could therefore not cause the observed difference. When cows were excluded, more litter accumulated in the vegetation. The formation of this litter layer may have added to the higher mineralization rates under vole grazing, through enhanced nutrient return through litter or through modification of microclimate. We conclude that different-sized herbivores have different effects on N cycling within the same habitat. Exclusion of large herbivores resulted in increased N annual mineralization under small herbivore grazing.     

Species-specific sprouting pattern in two dioecious Lindera shrubs: The role of physiological integration

Sprouting is recognized as an important genet persistence strategy for clonal woody plants, but the role of sprouting may differ between species and between sexes, depending on physiological integration. We tested the effect of physiological integration on the mortality, recruitment and growth of the sprouting male and female ramets of two closely related dioecious shrubs of Lindera, in a field experiment using girdling manipulation. Although between-sex differences observed were obscure, we found between-species differences in the sprouting patterns. The rates of ramet mortality and recruitment were significantly lower for L. praecox than L. triloba. In L. praecox genets, the ramet production was low, and the main ramets might actively translocate assimilates towards the small sprouted ramets, which then facilitates high ramet growth and survival (sprout-nursing strategy). Meanwhile, in L. triloba genets, although many ramets were recruited, assimilate translocation from the main ramets to the sprouted ramets might be less abundant, which causes high ramet mortality (sprout-turnover strategy). For a more general knowledge of the various sprouting strategies in clonal plants, our study demonstrated that inter-specific comparisons using girdling experiments at the whole-plant level could reveal the role of physiological integration on the link between the sprouting pattern and above-ground structures of clonal plants.     

Ramet dynamics in a centrifugally expanding clonal sedge: a matrix analysis

Carex humilis is a clonal sedge that can form distinct rings of densely aggregated ramets. We hypothesize that rings form because both production of new ramets and ramet dispersal are positively correlated to ramet size. This would lead to an overrepresentation of fast-moving and large ramets with high ramet production at the periphery, whereas slow-moving and small ramets with low ramet production would mainly be found in the interior of rings. We use matrix models to analyse how ramet populations both at the periphery and in the interior develop in the absence of ramet dispersal. We found that the stable size class distributions of ramets predicted by the models were not different from the distributions found in the field. Also, the asymptotic ramet population growth rates (λ₁) were the same. Hence, we conclude that rings would form even in the absence of a link between ramet dispersal and ramet production. Further analysis of the matrix models showed that the ramet population increases at the periphery but decreases in the interior of rings because medium and large ramets produce fewer large ramets in the interior than at the periphery. We also found that the temporal variance in λ₁ and transitions rates during the four study years was much higher at the periphery than in the interior. Our results suggest that rings may form because C. humilis ramets use below-ground resources from a much larger area than the one covered by the shoots. As the clone grows larger, the soil volume available to the ramets in the interior decreases because their access to soil outside the ring is cut-off by the ramets at the periphery. Ramet density in the interior is therefore decreasing.     

Contrasting effects of large herbivore grazing on smaller herbivores

Assemblages of large herbivores may compete for food or facilitate one another. However, small vertebrate herbivore species co-occurring with large herbivores may be affected by large herbivore grazing through changes in plant species composition, nutrient content and vegetation structure. These changes can be either positive or negative for the smaller herbivores, but this may depend on the species of small herbivores. We experimentally tested the impact of cattle grazing on habitat choice of European rabbits (Oryctolagus cuniculus) and common voles (Microtus arvalis). We excluded cattle for 7 years and measured changes in vegetation parameters, and the response of rabbits and voles. Rabbits were facilitated by cattle, whereas voles strongly preferred vegetation without cattle. The facilitation effect was stronger at low rabbit densities. Vegetation biomass and nitrogen concentration were not affected by cattle grazing, but vegetation height increased significantly where cattle were excluded. Plant species composition also changed following cattle exclusion; however, the main food plants of rabbits and voles remained abundant in each grazing treatment. We conclude that the response of both rabbits and voles predominantly reflect the differences in vegetation height in the presence and absence of cattle, but in a contrasting fashion. The difference in response between rabbits and voles may result from reduced perceived predation risk, which is lowest in high vegetation for voles, but in short vegetation for rabbits, which depend on their burrows for safety. The use of large herbivores in grassland conservation management can thus have a contrasting effect on different species of small herbivores.     

Demography of the clonal palm Geonoma brevispatha in a Neotropical swamp forest

Abstract We expected that in clonal plants with the phalanx growth form, prereproductive ramets function as a meristem bank that helps buffer ramet demography against environmental fluctuations. Specifically, we evaluated four expectations for the forest understorey clonal palm Geonoma brevispatha in a swamp forest in south‐eastern Brazil. Our expectations were are follows: (i) the vital rates of prereproductive ramets are more responsive to genet‐related characteristics (genet size, distance and size of the nearest genet neighbour, and reproductive ramet survivorship) than to environmental variables (canopy openness, basal area of small trees, flooding, and distance and size of the nearest large tree neighbour), whose effects are expected to affect ramet dynamics indirectly through physiological integration; (ii) the progress of prereproductive ramets to later stages is dependent on the death of reproductive ramets; (iii) the population dynamics of G. brevispatha is resistant to significant interannual changes in the vital rates of ramets, (i.e. the occurrence of good and bad years); and (iv) there are large negative covariances among demographic transitions within a given year, as well as between years, reflecting the flexibility of G. brevispatha to adjust to environmental variability. Expectations 1, 2 and 4 were confirmed, while expectation 3 was only partially confirmed. Matrix models predicted that the growth rate of the overall ramet population was significantly greater than unity in the first study year, but it was significantly smaller than 1.0 in the second study year. Ramet population growth rates at the level of individual genets were greater than unity in the majority of genets in the first year, but in the second year genets with growth rates smaller than unity prevailed. Ramet population dynamics seems to be resistant to the succession of good and bad years; in effect, the presence of a meristem bank of ramets helps stabilize λ.     

Clonal integration enhances flood tolerance of Spartina alterniflora daughter ramets

Recently, considerable attention has been paid to the invasion of the clonal plant Spartina alterniflora into coastal wetlands at lower elevations. In this experiment, we tested whether clonal integration improved flood tolerance in S. alterniflora daughter ramets. Daughter ramets at two growth stages (young and old ramets) were flooded to water levels of 0, 9 and 18 cm above the soil surface, and the rhizomes between mother and daughter ramets were either severed or left intact. Biomasses of connected ramets grown in controls or in shallow and deep water treatments were 119%, 108% and 149% higher in the old ramet group than those of severed ramets, respectively, whereas they were 3.0, 3.3 and 11.2 times higher in the young ramet group, respectively. At the end of the experimental period, the shoot height, connected with young ramets, in shallow and deep water treatments increased by 19% and 26%, respectively, over that in the control treatments, whereas the old ramets increased by 11% and 39%, respectively. In contrast, the shoot height of the severed young ramets was 27% and 26% lower in shallow and deep water treatments than in the control treatment, respectively. However, the shoot height of the severed old ramets remained constant with increasing water depth. We conclude that clonal integration enhances the flood tolerance of S. alterniflora daughter ramets, and the trait of clonal integration plays more important roles in severe flooding stress conditions and at early growth stages.     

Partial mechanical stimulation facilitates the growth of the rhizomatous plant Leymus secalinus: modulation by clonal integration

Background and Aims

Mechanical stimulation (MS) often induces plants to undergo thigmomorphogenesis and to synthesize an array of signalling substances. In clonal plants, connected ramets often share resources and hormones. However, little is known about whether and how clonal integration influences the ability of clonal plants to withstand MS. We hypothesized that the effects of MS may be modulated by clonal integration.

Methods

We conducted an experiment in which ramet pairs of Leymus secalinus were subjected to three treatments: (1) connected ramet pairs under a homogeneous condition [i.e. the proximal (relatively old) and distal (relatively young) ramets were not mechanically stressed]; (2) connected ramet pairs under a heterogeneous condition (i.e. the proximal ramet was mechanically stressed but the distal ramet was not); and (3) disconnected ramet pairs under the same condition as in treatment 2. At the end of the experiment, we harvested all plants and determined their biomass and allocation.

Key Results

Clonal integration had no significant influence on measured traits of distal L. secalinus ramets without MS. However, under MS, plants with distal ramets that were connected to a mother ramet produced more total plant biomass, below-ground biomass, ramets and total rhizome length than those that were not connected. Partial MS exerted local effects on stimulated ramets and remote effects on connected unstimulated ramets. Partial MS increased total biomass, root/shoot ratio, number of ramets and total rhizome length of stimulated proximal ramets, and increased total biomass, root weight ratio, number of ramets and total rhizome length of connected unstimulated ramets due to clonal integration.

Conclusions

These findings suggest that thigmomorphogenesis may protect plants from the stresses caused by high winds or trampling and that thigmomorphogenesis can be strongly modulated by the degree of clonal integration.     

Effects of inoculation with native arbuscular mycorrhizal fungi on clonal growth of Potentilla reptans and Fragaria moschata (Rosaceae)

Many clonal plants live in symbiosis with ubiquitous arbuscular mycorrhizal (AM) fungi, however, little is known about their interaction with respect to clonal reproduction and resource acquisition. The effects of arbuscular mycorrhiza on the growth and intraclonal integration between ramets of two stoloniferous species were studied experimentally in a nutritionally homogenous soil environment. Two species coexisting at the same field site, Potentilla reptans and Fragaria moschata, were selected as model plants for the study. Pairs of their ramets were grown in neighbouring pots with each ramet rooted separately. Four inoculation treatments were established: (1) both mother and daughter ramets remained non-inoculated, (2) both ramets were inoculated with a mixture of three native AM fungi from the site of plant origin, (3) only mother or (4) daughter ramet was inoculated. The stolons connecting the ramets were either left intact or were disrupted. Despite the consistent increase in phosphorus concentrations in inoculated plants, a negative growth response of both plant species to inoculation with AM fungi was observed and inoculated ramets produced fewer stolons and fewer offspring ramets and had lower total shoot dry weights as compared to non-inoculated ones. A difference in the extent of the negative mycorrhizal growth response was recorded between mother and daughter ramets of P. reptans, with daughter ramets being more susceptible. Due to AM effect on ramet performance, and thereby on the source-sink relationship, inoculation also significantly influenced biomass allocation within clonal fragments. Physiological integration between mother and daughter ramets was observed when their root systems were heterogeneous in terms of AM colonization. These results hence indicate the potential of mycorrhizal fungi to impact clonal growth traits of stoloniferous plant species, with possible consequences for their population dynamics.     

The significance of rhizome connection of semi-shrub Hedysarum laeve in an Inner Mongolian dune,China

Hedysarum laeve, a rhizomatous clonal semi-shrub, commonly dominates the inland dunes in semiarid areas of northern China. This species propagates vegetatively by extension of horizontal woody rhizomes resulting in programmed reiteration of apical and/or axillary meristems. In this study, the plants were experimentally manipulated by cutting rhizome connections and ¹⁴C-labelling techniques were employed to investigate the ecological significance of rhizome connections within the H. laeve clone. Severance of rhizome connections had a great effect on the performance of young ramets within a clone. Young ramets severed from their parent ramets experienced a significant reduction both in ramet growth and vegetative propagation, as compared with the intact young ramets. Within clonal fragments, consisting of three interconnected ramets including a mother ramet, a daughter ramet and a granddaughter ramet, ¹⁴C-photosynthates from the fed leaves of mother ramets were acropetally transported to all clonal component parts. The ¹⁴C-photosynthate translocation within the clonal fragment provides evidence that the young ramets were supported by their parent ramets. Our results suggest that the woody rhizome connections among the interconnected ramets are ecologically and strategically important for the species to grow in the sand dune habitat.     

Seasonal changes in response ofSenecio ovatus to grazing by the chrysomelid beetleChrysomela speciosissima

Summary The response ofSenecio ovatus to herbivory byChrysomela speciosissima (Coleoptera, Chrysomelidae) was studied in the Krušné hory Mountains, Czechoslovakia. The following questions were addressed: (1) what is the impact of natural levels of insect herbivory on the performance of individual plants, (2) how do the levels of herbivory change during the growing period, and (3) what is the relative importance of direct and indirect effects of herbivory? Ten experimental plots sprayed with metathion and ten controls were established. Damage to plants was quantified in the period of maximum herbivore activity (June) and again at the end of the growing season (August). In the first half of the season, total leaf area in grazed ramets was reduced by 30.3% compared to ungrazed ramets; half of this reduction was due to beetle consumption and half was due to indirect effects. Individual leaf area was not significantly different between treatments. Insecticide-treated ramets were taller and heavier than the controls. During the second half of the growing period the differences in ramet weight and leaf area disappeared because of compensatory growth. However, the insecticide-treated ramets remained taller and had higher seed numbers per capitulum. Neither number of capitula per ramet nor seed weight were significantly different between treatments. In the grazed population, the leaf area reduction (30.3%) in June resulted in 36.5% reduction in fecundity at the end of August.     

Environmentally-induced functional specialization for locally abundant resources in the stoloniferous herb Duchesnea indica

Reciprocally patchy environments, where the availability of two resources are patchily distributed and negatively correlated in each patch, are common in many ecosystems. Interconnected ramets of clonal plants can specialize in the uptake of locally abundant resources. Ramet pairs of the stoloniferous herb Duchesnea indica were grown in reciprocally patchy environments i.e., one ramet of a pair was grown in the high light but low water patch (high light patch) and the other in the low light but high water patch (high water patch). Biomass allocation pattern (root-shoot ratio), morphological traits (leaf area and root length) and physiological traits (photosynthetic rate and chlorophyll content) were altered in a way that potentially enables ramets to enhance the capture of the locally abundant resource (i.e., increase the capture of light resource in the high light patch and of water in the high water patch). As a result,biomass and number of ramets in the connected ramet pairs were greatly improved. Functional specialization of ramets, modified by clonal integration, may have contributed greatly to the growth increase of D. indica in the reciprocally patchy environment.     

Clonal plant allocation to daughter ramets is a simple function of parent size across species and nutrient levels

The evolution of clonal growth is a widespread phenomenon among plant species, characterized by the production of genetically identical clonal fragments (ramets) via rhizomes or stolons that form an interconnected clonal organism (genet). Clonal plant species are known to differ in their investment into ramet production, and exhibit considerable variation in ramet morphology both within and among species. While patterns of resource allocation are thought to be linked to a number of plant characteristics, many analyses are limited by uncertainty in how clonal plants determine the morphology and resources allocated to new ramets. In this study, we attempted to discern what aspects of parent ramets best predicted resource allocation to new daughter ramets, and the relationship between resource allocation and daughter ramet rhizome morphology. We grew two sedge species, Schoenoplectus tabernaemontani and Eleocharis elliptica, in a greenhouse under two levels of fertilizer addition. By harvesting daughter ramets that had initiated stem production, yet remained aphotosynthetic, we were able to isolate parental investment into non-independent daughter ramets at a point where daughter ramet spacer length became fixed. Our results indicate that parent ramets allocated a non-linear proportion of parent rhizome biomass to the production of daughter ramets. Moreover, this relationship was unaffected by environmental nutrient availability. Daughter ramet biomass, in turn, was strongly correlated with daughter ramet spacer length. These observations shed light on key processes governing clonal growth in plants, and their potential application in unifying allocational and morphological perspectives to explore the fitness implications of variability in clonal growth.     

外部干扰对根茎型克隆植物甘草自然种群植株生长及主要药用成分含量的影响

克隆植物, 特别是游击型克隆植物, 其分株在一定时间段往往通过连接结构(即间隔子)连接在一起形成克隆网络, 基于克隆网络的克隆植物特性显著提高了克隆植物应对外部干扰的能力。长期封育禁牧使得甘草(Glycyrrhiza uralensis)自然种群免受外部干扰, 却并没有促使受破坏的甘草自然种群快速恢复。因为克隆整合、克隆储存及克隆分株的选择性放置等克隆植物特性的存在, 适度的外部干扰有利于根茎型克隆植物甘草的生长与繁衍。但外部干扰与克隆植物特性在甘草种群恢复中的作用及其机理目前并不清楚。该研究以甘草自然种群为研究对象, 通过野外控制实验进行模拟动物采食和人工采挖干扰, 探讨不同程度干扰对甘草自然种群生长和主要药用成分含量的影响。结果表明在中度放牧采食干扰或者人工采挖干扰下, 甘草自然种群的分株密度、高度和地上生物量与对照相比均没有显著差异, 甘草表现出等补偿生长; 而在不同类型的重度干扰下, 甘草产生的响应不一致。重度放牧采食干扰下甘草表现出欠补偿生长, 而重度人工采挖干扰下甘草表现出超补偿生长。克隆植物特性在甘草应对外部干扰过程中发挥着重要作用。外部干扰一定程度上提高了甘草中甘草苷和甘草酸的含量, 从而提升了药材的品质。进一步的研究应该关注于甘草自然种群与环境因子之间的关系, 解析甘草自然种群退化的原因, 提出相应的复壮技术, 探讨可持续利用的管理模式。     

Responses of rhizomatous grass Phragmites communis to wind erosion: effects on biomass allocation

Background and aims

The ability of modifying biomass allocation to deal with different environmental stress is an important mechanism for plant population expansion and maintenance in the unstable dune environment where wind erosion persists. However, how biomass is partitioned between horizontal rhizome extension and vertical ramet growth in response to wind erosion has not been fully understood. The objective of this study was to explore how wind erosion affected the relationship between horizontal rhizome extension and vertical ramet growth using a common rhizomatous perennial grass, Phragmites communis.

Methods

We dug 300 cm?×?200 cm, 80 cm deep pits in a garden experiment plot. Clonal fragments of P. communis were planted individually at a depth of 40 cm in these pits for 4 weeks before treatments. Surface sand was gradually removed to the final depth of 0 (control), 10, 20, 30 and 40 cm (maximum sand removal). Ramet emergence time, rhizome-based and tiller-based ramet number, rhizome number and length, biomass of vertically and horizontally oriented structures were monitored at the end of the experiment.

Results

With increasing erosion depth, the proportion of tiller-based ramets (in total number of ramets) increased, whereas that of rhizome-based ramets decreased. With increasing erosion depth, the percentage of vertically oriented structures biomass in total biomass increased significantly, whereas that of horizontally oriented structures biomass decreased.

Conclusions

The changes in biomass allocation (i.e., more allocation in vertical than horizontal biomass) together with a trade-off in tiller-based and rhizome-based ramets may enable P. communis to make better use of the resources in erosion conditions and maximize plant population expansion and maintenance.     

Integration in the clonal plant <Emphasis Type="Italic">Eriophorum angustifolium</Emphasis>: an experiment with a three-member-clonal system in a patchy environment

A clonal plant in heterogeneous environments is usually expected to profit from resource exchange via a clonal network where ramets placed in contrasting environments can specialise so to acquire the most abundant resources. An experiment was designed using the three member clonal system of Eriophorum angustifolium, which consisted of one parent ramet growing in a resource poor environment and two offspring: one was limited in growth by nutrients while the other was light limited; the contrast in availability of limited resources between the offspring ramets was high, medium or none, with the system either connected or severed. The total resource availability was the same in all treatments. We proposed four possible scenarios for the system: offspring ramets will share resources via the deficient parent ramet, and the whole clone will profit from the contrasting environment (scenario 1); offspring ramets will support exclusively the parent ramet, and the whole clone will profit from a homogeneous environment (scenario 2); offspring ramets will stop the export of the limiting resource to the parent ramet, with split and connected treatments not differing (scenario 3); and offspring ramets will exhaust the carbon stored in the biomass of the parental ramet; offspring ramet will profit from connection (scenario 4). In the experiment, the limiting resources were sent to the strongest sink (scenario 2). The parent ramet growing in a deficient environment received the highest support in the treatment where both offspring ramets were growing in the same conditions (no-contrast treatment). Production of new shoots, but not biomass of whole clone, was supported in a homogenous environment. The experiment revealed that multiple stresses might prohibit free exchange of limiting resources via the clonal network and supports the idea that experimental studies on more complex clones are essential for understanding the costs and benefits of clonal growth.     

Environmentally-induced functional specialization for locally abundant resources in the stoloniferous herb Duchesnea indica

Reciprocally patchy environments, where the availability of two resources are patchily distributed and negatively correlated in each patch, are common in many ecosystems. Interconnected ramets of clonal plants can specialize in the uptake of locally abundant resources. Ramet pairs of the stoloniferous herb Duchesnea indica were grown in reciprocally patchy environments i.e., one ramet of a pair was grown in the high light but low water patch (high light patch) and the other in the low light but high water patch (high water patch). Biomass allocation pattern (root-shoot ratio), morphological traits (leaf area and root length) and physiological traits (photosynthetic rate and chlorophyll content) were altered in a way that potentially enables ramets to enhance the capture of the locally abundant resource (i.e., increase the capture of light resource in the high light patch and of water in the high water patch). As a result,biomass and number of ramets in the connected ramet pairs were greatly improved. Functional specialization of ramets, modified by clonal integration, may have contributed greatly to the growth increase of D. indica in the reciprocally patchy environment.     

Sexual reproduction and clonal growth in Reinhardtia gracilis (Palmae), an understory tropical palm

Patterns of sexual reproduction and clonal growth were investigated in the understory palm Reinhardtia gracilis var. gracilior over a 3-yr period. R. gracilis is a very abundant clonal palm in the tropical rain forest of Los Tuxtlas, Veracruz, México. Because ramets form clumps, genets are easily identified in the field. Genets were monitored in a 0.5-ha area, and classified by size according to the number of ramets they possessed. In contrast to clonal growth, sexual reproduction was highly dependent on genet size. The probability of reproduction, the number of inflorescences, and the number of fruits produced were positively correlated with genet size. However, neither the probability of producing a ramet, nor the number of ramets produced per genet were correlated with genet size. Over the 3 yr of study, 55% of the genet population had at least one ramet with reproductive structures, while <1% (a single genet in one year) had six ramets with flowers. Thirty-two percent of the mature genets reproduced during each of three consecutive years. In contrast, 58% of the genets produced no new ramets during these 3 yr. No evidence was found of a trade-off between clonal growth and sexual reproduction. Ramet production increases genet size and this in turn increases genet reproductive performance. Clonal growth in this species may be viewed as a growth strategy that tends to maximize genet fitness.     

Clonal integration affects growth and sediment properties of the first ramet generation,but not later ramet generations under severe light stress

克隆整合影响严重光胁迫下第一分株世代的生长和沉积物特征但不影响 后续分株世代的生长和沉积物特征 克隆整合通过缓冲环境压力和提高资源获取效率使克隆植物受益。然而,在一个克隆系统中,受益于克隆整合的连接分株世代的数量很少受到关注。我们进行了一个盆栽实验来评估沉水植物苦草 (Vallisneria natans)克隆系统内的生理整合程度,该克隆系统由一个母株和3个依次连接的后代分株组成。 母株生长在正常光照下,而后代分株被严重遮荫。母株与后代分株间的匍匐茎被切断或保持连接,但3个后代分株之间的连接仍然存在。与遮荫的后代分株连接时,苦草未遮荫的母株的光合能力显著增强,但其生物量积累大大减少。克隆整合显著增加了第一分株世代(相邻分株)的生物量积累和土壤的碳氮可用性、胞外酶活性和微生物生物量,但没有增加后续分株世代的这些特征。我们的结果表明,在严重光胁迫下,来自苦草母株的支持可能仅限于克隆系统中相邻的后代分株,这暗示着一个分株世代的效应。我们的结果有助于更好地理解克隆植物的层次结构和分段化。这些发现表明克隆整合程度在分株种群的生态相互作用中起着至关重要的作用。