No long-term costs of meristem allocation to flowering in stoloniferous Trifolium species

Clonal plants propagate by means of clonal growth and sexual reproduction. The commitment of meristems to branching and flowering govern the expression of these two mutually exclusive life-history functions. We used a modelling and an experimental approach to examine the consequences of a structural trade-off between flowering and clonal growth on future growth and fitness in stoloniferous species with a determinate module architecture. The model revealed negative effects of flowering on vegetative growth due to a structural trade-off at the meristem level. Total fecundity was maximized at intermediate flowering frequencies. In addition, optimal meristem commitment to flowering depended strongly on the time available for growth and reproduction. This indicates an interaction between optimal flowering frequency, the length of the growing period and the rate of ontogenetic development. The greenhouse study made use of 15 genotypes of two closely related, stoloniferous Trifolium species. Despite the existence of a structural trade-off at the meristem level, we found no evidence for costs of flowering on the whole-plant level. High allocation to flowering did not result in reduced plant performance (biomass and module production) and total fecundity, indicating that there were no demographic costs of meristem investment to different life-history functions. Flowering frequencies never exceeded the model prediction for optimal commitment of meristems to sexual reproduction, suggesting strong past selection to eliminate high levels of meristem allocation to flowering. Hence, clonal growth seems to have evolutionary priority over sexual reproduction in our species. This revised version was published online in November 2006 with corrections to the Cover Date.     

Life history trade-offs in Amphibromus scabrivalvis (Poaceae): allocation to clonal growth,storage, and cleistogamous reproduction

Life history trade-offs among clonal growth, storage, and sexual reproduction were investigated in the perennial grass Amphibromus scabrivalvis in relation to soil nutrients. This species exhibits clonal growth by producing rhizomes and stores reserves in the form of basal corms; seeds are matured in cleistogamous spikelets on panicles enclosed within the leaf sheaths along each culm. Ten seed-derived genotypes (clones) were separated into 72 ramets and planted in the greenhouse. Control ramets received only water while the remainder received fertilizer applied every 2 wk. Twenty-four ramets were harvested per clone at 11, 20, and 26 wk. The dry mass of corms, rhizomes, roots, shoots, and seeds were recorded. Biomass partitioning to rhizomes provided a measure of carbon allocation to clonal growth, partitioning to corms provided a measure of allocation to storage, and partitioning to seeds provided a measure of allocation to sexual reproduction. Allocation to most organs was significantly influenced by clone identity; fertilizer significantly increased allocation to corms and seeds at 20 wk, but never affected rhizome allocation at any age. Corm allocation increased from 2% at 11 wk to 27% at 26 wk; rhizome allocation decreased from 10% at 11 wk to 3% at 26 wk. Significant negative relationships were detected for rhizome vs. seed and corm vs. rhizome allocation in fertilized clones at 20 wk. This suggests an age-dependent physiological life history trade-off between clonality and sexual reproduction and between clonality and storage. In contrast, a significant positive relationship was consistently noted for corm vs. seed allocation in fertilized and unfertilized clones at 20 and 26 wk. The absence of a trade-off between storage and sexual reproduction may indicate that these two processes are not necessarily mutually exclusive components of life history.     

Effects of light on the growth and clonal reproduction of Ligularia virgaurea

Ligularia virgaurea is a perennial herb that is widely distributed in the alpine meadow on the eastern Qinghai-Tibet plateau.We investigated the patterns of growth and reproduction of L.virgaurea under two contrasting levels of light conditions for two continuous growing seasons.Our results showed that the light affects on the maximum relative growth rate,the shoot weight ratio and the root weight ratio differed between the two growing seasons.L.virgaurea reproduced initially through rhizome in the second growing season,rather than sexual reproduction.The proportion of genets with clonal reproduction decreased under shaded conditions.A minimum genet size should be attained for clonal reproduction to begin under the shaded conditions.There was a positive linear relationship between clonal reproduction and genet size.Light level affected the allocation of total biomass to clonal structures,with less allocation under the full natural irradiance than under the shaded conditions.There seemed to be a trade-off between vegetative growth and clonal reproduction under the full natural irradiance,in terms of smaller relative growth rates of genets with clonal reproduction than those without clonal reproduction.L.virgaurea emphasized clonal reproduction under the full natural irradiance,while the plant emphasized vegetative growth under the shaded conditions.     

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.     

Heavy metals modify costs of reproduction and clonal growth in the stoloniferous herb Potentilla anserina

We examined costs of sexual reproduction and clonal propagation, and their consequences for resource allocation in the clonal stoloniferous herb, Potentilla anserina, a typical pioneer species in disturbed areas. We used heavy-metal treatment in soil to create unfavourable growing conditions, because costs of reproduction are more likely to be expressed when resources are limited. We also studied whether heavy metals affect the plasticity of clonal growth form that enables the plants to avoid poor growing conditions. Ramets collected from field were grown in a greenhouse under the heavy-metal treatment consisting of a control and two levels of heavy-metals added in soil. Clonal propagation was costly in terms of total biomass of flowering ramets. Also the costs of sexual reproduction were detected in flowering ramets. Contrary to our predictions, the costs of flower production were visible in the control but not in the heavy-metal contaminated plants. Only the flowering ramets were able to produce longer stolons under heavy-metal stress, but the stolon biomass was not affected by heavy metals. Results indicate that clonal propagation and sexual reproduction may be costly for P. anserina. However, the costs are modified by heavy-metal contamination.Co-ordinating editor: J. Tuomi     

Density and the commitment of apical meristems to clonal growth and reproduction in Hieracium pilosella

Summary We examined responses to population density in the commitment of apical meristems to reproduction and clonal growth in a rosette-forming, stoloniferous herb (Hieracium pilosella). Despite close physiological coupling between the evocation of the terminal inflorescence bud and the development of one or more axillary buds into stolons, the allocation of meristems was extremely plastic.Genets at the higher sowing densities showed density-dependent mortality consistent with self-thinning along a-3/2 trajectory. The probability of inflorescence evocation and associated stolon development was negatively dependent on surviving density. The proportinal distribution of primary stolons amongst genets became strikingly more unequal (expressed as the Gini coefficient) with increasing density. Clonal growth was resolved into the number of primary stolons per stoloniferous genet and the extent of stolon branching (i.e. number of apices per primary stolon); both showed strongly negative density-dependence. Reproduction, expressed as the mean number of flowering capitula per stoloniferous genet, declined 15-fold with increasing density; although theoretically expected to be unity, greater values resulted from capitulum production by attached secondary rosettes and lower values reflected the increasing abortion rate of inflorescence buds with increasing density.Both the total number of apices produced per unit area and the corresponding number of reproductive apices were maximal at intermediate surviving densities (700–1,000 m^-2). The balance between reproductive and clonal growth may be expressed as the probability of an apical meristem producing a capitulum, that also peaked sharply at intermediate density. This finding does not conform with linear models that predict a shift from vegetative growth to sexual reproduction with increasing population density.     

The effect of pollination on resource allocation among sexual reproduction,clonal reproduction,and vegetative growth in <Emphasis Type="Italic">Sagittaria potamogetifolia</Emphasis> (Alismataceae)

In order to evaluate the effects of pollination on resource allocation in the marsh herb Sagittaria potamogetifolia, experimental manipulation of pollination efficiency on the pattern of resource allocation was accessed by the proportion of dry weight measurements of sexual, vegetative, and clonal organs. In trials where half of the flowers were pollinated, a significant increase of resource allocated to sexual production and decrease to vegetative production resulted compared to plants that received no pollination. In trials where pollination was 100%, these two reproductive components showed the same trend, but less dramatically. This may support the idea that the trade-offs would be more pronounced when the resource was scarce. Besides, a higher inflorescence production with a lower fruit reproduction occurred as a consequence of decreased pollination level. This increased inflorescence production may be a mechanism to promote outcrossing by enhancing floral attraction or by synchronizing reproductive activity with insect pollinators. Examination for possible trade-offs in resource allocation revealed that there was also a trade-off caused by pollination between fruits plus flowers and bulbils production, which might have detrimental effects on the survival of individuals and populations, but promotes outcrossing and genetic variability.     

不同高程短尖苔草对水位变化的生长及繁殖响应

在淡水湿地生态系统中,水位通常是制约植被生长和繁殖动态的关键因素,进而对物种组成、群落演替和植被分布格局产生决定性影响。无性繁殖是洞庭湖湿地克隆植物适应环境胁迫的重要策略之一,以洞庭湖湿地典型克隆植物-短尖苔草(Carex brevicuspis C.B.Clarke)为对象,研究了不同分布高程(23.7 m和25.8 m)的植物对水位变化(0 cm,-15 cm,-30 cm)的生长和繁殖特征响应。结果表明:水位变化对不同分布高程分布短尖苔草的生长和克隆繁殖特征均产生显著影响(P0.05)。对高程区的短尖苔草而言,克隆繁殖特征如分株数、分株总生物量、芽数和芽生物量随水位的降低而增加,而对生长特征(株高及总生物量)无显著影响(P0.05),表明适当干旱有利于高程区苔草的克隆繁殖。对于低程区分布短尖苔草而言,水位变化对其生长特征有显著影响(P0.05),如株高和总生物量随着水位的降低而增加;分株数和总芽生物量等克隆繁殖特征则随水位的降低而减少,而水位对低程区短尖苔草的分株总生物量和总芽数影响不显著(P0.05)。因此,短尖苔草的克隆繁殖特征除受到水位的影响外,还受其分布高程的影响。可见,同一种短尖苔草因长期适应于不同生境而对相同的环境胁迫表现出了不同的生长繁殖策略,     

Significance of the simultaneous growth of vegetative and reproductive organs in the prostrate annual Chamaesyce maculata (L.) Small (Euphorbiaceae)

To elucidate the significance of the simultaneous growth of vegetative and reproductive organs in the prostrate annual Chamaesyce maculata (L.) Small (Euphorbiaceae) from the standpoint of meristem allocation, we investigated plant architecture, meristem allocation, and the spatial and temporal patterns in vegetative growth and reproduction in the reproductive stage. The numbers of secondary and tertiary shoots successively increased by branching in the reproductive stage, and the sum of shoot length was greater in secondary shoots than in primary shoots. The specific shoot length (shoot length per shoot biomass) was greater in lateral shoots than in primary shoots, indicating efficient lateral shoot elongation. The internode length was shorter in secondary shoots than in primary shoots, increasing the number of nodes per shoot length in secondary shoots. Many nodes on a shoot generated two meristems, one of which committed to a flower and one to a lateral shoot. The number of reproductive meristems was greatest in tertiary shoots, and 96% of total reproductive meristems on shoots were generated in lateral shoots. On almost all nodes, the reproductive meristem developed into a flower, and 95–98% of the flowers produced a fruit. Therefore, vegetative growth by branching in the reproductive stage contributed to the increase in reproductive outputs. From the standpoint of meristem allocation, the simultaneous growth of vegetative and reproductive organs in prostrate plant species might be important for increasing the number of growth and reproductive meristems, resulting in the increase in reproductive outputs.     

The relative importance of sexual and clonal reproduction for population growth in the perennial herb Fragaria vesca

The relative importance of sexual and clonal reproduction for population growth in clonal plants is highly variable. Clonal reproduction is often more important than sexual reproduction but there is considerable interspecific variation and the importance of the two reproductive modes can change with environmental conditions. We carried out a demographic study on the woodland strawberry (Fragaria vesca), a widespread clonal herb, at 12 sites in Switzerland during 2 years. Study sites were selected in two different habitats, i.e., forest and forest edge. We used periodic matrix models to estimate annual population growth rates and carried out prospective analyses to identify life cycle components that influence population growth rates most. Retrospective analyses were applied to study how the two different habitats affected population dynamics. Furthermore, we tested whether trade-offs between sexual and clonal reproduction occurred. There were large differences in annual population growth rates between sites and large within-site differences between years. Results of the prospective analyses clearly indicate that clonal reproduction is the dominant reproductive pathway whereas sexual reproduction is rather insignificant for population growth. Compared to forest habitats, forest edge habitats had higher population growth rates in the first year but smaller growth rates in the second year. We attribute these differing habitat effects to different water availabilities during consecutive years. No trade-offs between sexual and clonal reproduction were found. In conclusion, population growth of F. vesca relies heavily on clonal reproduction. Furthermore, reproduction and survival rates of F. vesca depend highly on spatio-temporal variation of environmental conditions.     

Stress promotes changes in resource allocation to growth and reproduction in a simultaneous hermaphrodite with indeterminate growth

In iteroparous animals, investment in growth is compromised by investment in reproduction, especially in species with indeterminate growth. Life‐history theory predicts that growth should be favoured over reproduction, assuming size‐related fecundity or survival. Hence, increase body condition represents an increase in reproductive potential. Simultaneous hermaphrodites should adjust their resource allocation to each sex function in response to current conditions but, recently, it has been suggested that, in hermaphrodites, gender allocation should be considered as a three‐way trade‐off, including the investment in somatic growth. Due to the higher costs involved, the female function is affected to a greater extent by environmentally stressful conditions rather than the male function. To examine this, we induced stress in the hermaphroditic earthworm Eisenia fetida (Savigny, 1826) and looked for changes in resource allocation in nonreproductive and reproductive individuals. Experimental stress was induced by using tweezers to elicit contractile escape movements. We predicted that stressed earthworms would preferentially allocate resources to growth. In nonreproductive individuals, however, stress had a negative effect on growth, although weight recovery was rapid once manipulation ceased, indicating the importance of body condition, as well as the existence of mechanisms of compensatory growth for growth trajectories in this earthworm species. The response of reproductive individuals was consistent with our expectation: (1) stressed worms maintained their growth rate at the expense of current reproduction and (2) stressed earthworms laid 25% fewer cocoons, which were 30% lighter than cocoons laid by control earthworms. The present results suggest that E. fetida regulates its reproductive effort and that future reproduction has more impact on its fitness than current reproduction. The trade‐off between current and future reproduction should be taken into consideration in models of sex allocation in simultaneous hermaphrodites. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 91 , 593–600.     

Sexual reproduction, clonal diversity and genetic differentiation in patchily distributed populations of the temperate forest herb Paris quadrifolia (Trilliaceae)

Clonal plant species have been shown to adopt different strategies to persist in heterogeneous environments by changing relative investments in sexual reproduction and clonal propagation. As a result, clonal diversity and genetic variation may be different along environmental gradients. We examined the regional and local population structure of the clonal rhizomatous forest herb Paris quadrifolia in a complex of forest fragments in Voeren (Belgium). Relationships between population size (the number of shoots), shoot density (the number of shoots per m²) and local growth conditions were investigated for 47 populations. Clonal diversity and genetic variation within and among 19 populations were investigated using amplified fragment length polymorphism markers. To assess the importance of sexual reproduction, seed set, seed weight and germination success were determined in 18 populations. As predicted, local growth conditions largely affected population distribution, size and density of P. quadrifolia. Populations occurring in moist and relatively productive sites contained significantly more shoots. Here, shoots were also much more sparsely distributed compared to populations occurring in dry and relatively unproductive sites, where shoots showed a strongly aggregated distribution pattern. Clonal diversity was relatively high, compared with other clonal species (G/N ratio = 0.43 and Simpson’s D=0.81). Clonal diversity significantly (P<0.01) decreased with increasing shoot density while molecular genetic variation was significantly (P<0.01) affected by population size and local environmental conditions. Lack of recruitment and out-competition of less-adapted genotypes may explain the decreased genetic variation in dry sites. Analysis of molecular variance revealed significant genetic variation among populations (Φ _ST=0.42, P<0.001), whereas pairwise genetic distances were not correlated to geographic distances, suggesting that gene flow among populations is limited. Finally, the number of generative shoots, the number of seeds per fruit and seed weight were significantly and positively related to population size and local growth conditions. We conclude that under stressful conditions populations of clonal forest plant species can slowly evolve into remnant populations characterized by low levels of genetic variation and limited sexual reproduction. Conservation of suitable habitat conditions is therefore a prerequisite for effective long-term conservation of clonal forest plant species.     

Interactions in the patterns of vegetative growth and reproduction in woody dioecious plants

Summary Interactions between vegetative growth and reproduction were evaluated in Peumus boldus, Lithraea caustica and Laretia acaulis, three woody dioecious species in central Chile. Phenological observations were made periodically on marked branches of male and female plants, and biomass allocation (dry weight) to vegetative and reproductive tissues was measured. The magnitude of flowering was evaluated in groups of plants in three successive seasons. The patterns of activities are species- and sex-dependent, and cycles of 2–4 years have been established. Branches that produce flowers either do not grow or grow less than branches without flowers, and males and females have differential resource allocation: male branches attain higher biomass values. Groups of plants show seasonal behavior that suggest synchrony in their reproductive activities.     

A change from phalanx to guerrilla growth form is an effective strategy to acclimate to sedimentation in a wetland sedge species Carex brevicuspis (Cyperaceae)

The rhizomatous sedge Carex brevicuspis can produce clumping ramets from shortened rhizomes (phalanx) and spreading ramets from elongated rhizomes (guerrilla) to form a combined clonal growth form. In this paper, changes in clonal growth and biomass allocation pattern of C. brevicuspis in response to sedimentation were studied. Four sedimentation depths (0, 3, 6, and 9 cm) were applied to 48 ramets in a randomized block design. Plants were harvested after 20 weeks. With increasing sedimentation depth, the proportion of spreading ramets to total ramets increased from 19.6% in 0 cm to 92.9% in 9 cm sedimentation treatments, whereas that of clumping ramets decreased from 80.4% to 7.1%, indicating a change of clonal growth form from phalanx to guerrilla as a response to sedimentation. With increasing sedimentation depth, biomass allocation to shoots and roots did not change, but rhizome mass ratio increased from 2.7% in 0 cm to 7.2% in 9 cm sedimentation treatments, suggesting that production of long rhizomes changes biomass allocation pattern. The results show that plasticity of clonal growth forms, by which more spreading ramets are produced, is an effective strategy to avoid sedimentation stress under our experimental conditions.     

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。     

Trade-offs among growth, clonal, and sexual reproduction in an invasive plant Spartina alterniflora responding to inundation and clonal integration

The purpose of this article was to study the trade-offs among vegetative growth, clonal, and sexual reproduction in an aquatic invasive weed Spartina alterniflora that experienced different inundation depths and clonal integration. Here, the rhizome connections between mother and daughter ramets were either severed or left intact. Subsequently, these clones were flooded with water levels of 0, 9, and 18 cm above the soil surface. Severing rhizomes decreased growth and clonal reproduction of daughter ramets, and increased those of mother ramets grown in shallow and deep water. The daughter ramets disconnected from mother ramets did not flower, while sexual reproduction of mother ramets was not affected by severing. Clonal integration only benefited the total rhizome length, rhizome biomass, and number of rhizomes of the whole clones in non-inundation conditions. Furthermore, growth and clonal reproduction of mother, daughter ramets, and the whole clone decreased with inundation depth, whereas sexual reproduction of mother ramets and the whole clones increased. We concluded that the trade-offs among growth, clonal, and sexual reproduction of S. alterniflora would be affected by inundation depth, but not by clonal integration.     

THE COST OF MERISTEM LIMITATION IN POLYGONUM ARENASTRUM: NEGATIVE GENETIC CORRELATIONS BETWEEN FECUNDITY AND GROWTH

Growth and reproduction in higher plants depend on meristems, which have three developmental fates. A meristem can become reproductive, but doing so terminates its activity, it can differentiate vegetatively, or it can remain quiescent for extended periods. The first two fates are mutually exclusive, and only the second leads to the production of additional meristems for subsequent growth and reproduction. In Polygonum arenastrum (frequently referred to as P. aviculare in North American Floras), an annual species lacking quiescent meristems, a quantitative genetic analysis of inbred full-sibling families revealed genetic variation in the developmental pattern of axillary meristem commitment to vegetative growth versus reproduction. Developmental variation resulted in family differences in the age of first reproduction, in age-specific fecundity and growth, and in final plant size and reproductive output. Furthermore, there were strong negative genetic correlations between age-specific growth and fecundity. Early commitment of meristems to reproduction favors high early fecundity, but reduces the number of meristems available for vegetative differentiation, and leads to lowered growth rates and fecundity later in life, when meristems are limiting. Conversely, meristem commitment to vegetative growth early in life results in low early fecundity but high late fecundity and growth. Meristem limitation, like resource limitation, is a proximate mechanism that generates trade-offs between life history traits. Differences between meristem limitation and resource limitation are discussed. Meristem limitation leads automatically to a senescent life history because of the determinate fate of reproductive meristems. Developmental characters were also found to be genetically correlated with metamer characters (leaf size, internode length) and seed size in this selfing species. The pattern of correlation is suggestive of selection for particular suites of life history and morphological characters.     

Water availability and population origin affect the expression of the tradeoff between reproduction and growth in Plantago coronopus

Investment in reproduction and growth represent a classic tradeoff with implication for life history evolution. The local environment can play a major role in the magnitude and evolutionary consequences of such a tradeoff. Here, we examined the investment in reproductive and vegetative tissue in 40 maternal half‐sib families from four different populations of the herb Plantago coronopus growing in either a dry or wet greenhouse environment. Plants originated from populations with an annual or a perennial life form, with annuals prevailing in drier habitats with greater seasonal variation in both temperature and precipitation. We found that water availability affected the expression of the tradeoff (both phenotypic and genetic) between reproduction and growth, being most accentuated under dry condition. However, populations responded very differently to water treatments. Plants from annual populations showed a similar response to drought condition with little variation among maternal families, suggesting a history of selection favouring genotypes with high allocation to reproduction when water availability is low. Plants from annual populations also expressed the highest level of plasticity. For the perennial populations, one showed a large variation among maternal families in resource allocation and expressed significant negative genetic correlations between reproductive and vegetative biomass under drought. The other perennial population showed less variation in response to treatment and had trait values similar to those of the annuals, although it was significantly less plastic. We stress the importance of considering intraspecific variation in response to environmental change such as drought, as conspecific plants exhibited very different abilities and strategies to respond to high versus low water availability even among geographically close populations.     

Differential effects of water depth and sediment type on clonal growth of the submersed macrophyte <Emphasis Type="Italic">Vallisneria natans</Emphasis>

The response of clonal growth and ramet morphology to water depth (from 60 to 260 cm) and sediment type (sand versus organic clay) was investigated for the stoloniferous submersed macrophyte Vallisneria natans in an outdoor pond experiment. Results showed that water depth significantly affected clonal growth of V. natans in terms of clone weight, number of ramets, number of generations, clonal radius and stolon length. V. natans showed an optimal clonal growth at water depths of 110–160 cm, but at greater depths clonal growth was severely retarded. A high allometric effect was exhibited in ramet morphology. Along the sequentially produced ramet generations, ramet weight and plant height decreased while stolon length and the root:leaf weight ratio increased. When using ramet generations as covariate, sediment type rather than water depth more strongly affected the ramet characteristics. For plants grown in clay, ramet weight, ramet height and stolon length were greater, and plants exhibited lower root:leaf weight ratio. These data suggest that water depth and sediment type have differential effects on clonal growth of V. natans: Water depth appears primarily to affect numerical increase in ramets and spatial spread, whereas sediment type mainly affects biomass accumulation and biomass allocation. Handling editor: S. M. Thomaz     

Effects of ramet clipping and nutrient availability on growth and biomass allocation of yellow nutsedge

A glasshouse experiment was conducted to examine how the interactions of nutrient availability and partial ramet clipping affect growth, reproduction and biomass allocation of Cyperus esculentus, an invasive sedge. The plants sprouting from tubers were grown at low and high nutrient levels, and were subject either to no clipping, one, two or three clippings, with each clipping cutting half of the existing ramets at soil level. Our results show that nutrient availability and clipping frequency tended to independently affect most of growth, reproduction and biomass allocation parameters of Cyperus esculentus examined in the present study. Increased supply of nutrients led to an increase in plant productivity and its associated traits. All of the traits, except for the number of ramets, displayed a decreasing pattern with increasing clipping frequency, indicating that Cyperus esculentus had undercompensatory responses to ramet clipping. It is likely that the patterns of plants response to clipping are species specific, and depend on morphological characters of species. Its susceptibility to ramet clipping can offer opportunities for controlling this invasive species through mechanical methods such as mowing. Clipping had little effects on biomass allocation; however, root weight fraction increased with increasing clipping frequency. While nutrient availability and clipping frequency had no influence on leaf carbon concentration at harvest, both of them increased leaf nitrogen concentration, and hence reduced leaf C/N ratio.