斑苦竹无性系种群克隆生长格局动态的研究

采用“例逐龄级累加法”（ＲＡＡ）研究了缙云山斑苦竹无性系种群的克隆生长格局动态,以及无性系分株克隆生长型的动态趋势．结果表明,作为复轴型的斑苦竹,其无性系种群随时间进程表现为聚集程度逐渐降低的集群分布格局．在自然条件下,斑苦竹更多地表现出单轴型的繁殖趋势．应用ＲＡＡ分析植物种群,尤其是竹类植物种群前期的克隆生长格局的动态,结果可靠,具有重要的应用价值．     

Morphological responses to nutrient availability in four clonal herbs

This paper examines morphological plasticity of clonal plants of contrasting habitats and of contrasting architectures in response to nutrient supply. The hypotheses were tested that plants from rich habitats possess greater plasticity in response to variation in resource supply than species from poor habitats, and that rhizomatous species are less plastic in their response than stoloniferous species. Two sympodial rhizomatous herbs (Carex flacca, C. hirta) and two monopodial stoloniferous herbs (Trifolium fragiferum, T. repens) were subjected to four levels of nutrient supply in a garden experiment. One of the two species of each genus (C. hirta, T. repens) is from fertile and the other from infertile habitats. We measured 1) whole plant characters: total plant dry weight, number of modules (product of a single apical meristem) and number of ramets; 2) ramet characters: ramet leaf area and ramet height; and 3) spacer characters: branches per module, length per module and length per module internode.All measured characters in the Trifolium species significantly responded to treatment: the values for all measured characters increased with higher levels of fertilization. The differences in plant characters between fertilization levels were larger in Trifolium repens than in T. fragiferum in terms of whole plant characters, ramet characters and stolon internode length. The two Carex species did not differ in their responses to treatment in terms of most characters measured. In ramet characters and in some whole plant characters the species from fertile habitats were more plastic than those from infertile habitats. In spacer characters this pattern was not found. Foraging could not be demonstrated unequivocally.Morphological plasticity in the stoloniferous (Trifolium) species was much larger than in the rhizomatous (Carex) species. This seems in accordance with a foremost storage function of rhizomes, as against a foremost explorative function of stolons.     

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     

Field evidence of plastic growth responses to habitat heterogeneity in the clonal herbRanunculus repens

Stolon internode lengths were measured on plants of the clonal herbRanunculus repens growing in a hay meadow which was subject to disturbance by mole (Talpa europaea) activity. Within the site three habitat types were recognized: closed grassland, the open ground of fresh molehills and the grass-molehill boundary. The lengths of stolon internodes ofR. repens differed significantly in each of the three habitats. The shortest internodes occurred on stolons on the open molchills. The longest occurred in the closed grassland habitat. The type of habitat in which parent ramets were rooted did not significantly influence the length of internodes on their daughter stolons. The length of a stolon internode was determined by its immediate surrounding habitat type. Consecutive internode lengths on a given stolon showed considerable plasticity, shortening significantly as stolons spread onto molehills from surrounding habitats, and increasing significantly as stolons advanced from a molehill into the surrounding closed grassland habitat. These results are consistent with the proposition that under favorable conditions (on the molehills, where resources are expected to be more abundant, and competition absent) internode lengths shorten and the plant forages intensively, whereas under conditions of low resource availability (in the closed grassland, where competition occurs) internode lengths increase, allowingR. repens to forage extensively. Such morphological plasticity may promote more efficient exploitation of resource-rich sites and more rapid vacation of resource-poor sites.     

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.     

Differential response to shading in orthotropic and plagiotropic shoots of the clonal herb Glechoma hirsuta

Previous studies have shown that internodes and petioles of closely related erect and stoloniferous species show marked differences in their response to shading. Vertical structures show strong elongation responses while horizontal structures show significantly smaller elongation responses. This paper reports an experiment designed to test whether internodes and petioles on orthotropic (vertically oriented) and plagiotropic (horizontally oriented) shoots produced by the same plant, also show such differential responses. The study species, Glechoma hirsuta, produces plagiotropic shoots in its vegetative stage and orthotropic shoots during its generative stage. Shoots of G. hirsuta were grown either in full daylight or under simulated canopy shade. Internode and petiole elongation, biomass investment patterns and other growth-related parameters were measured on plants in each light treatment. In orthotropic shoots the length of internodes responded more strongly to shading than the length of petioles, while the opposite was true for plagiotropic shoots, confirming the hypothesis that vertical spacers have higher degrees of shade-induced plasticity than horizontally oriented spacers. Growth and development of horizontally oriented shoots was reduced by shading, whereas that of vertically oriented shoots was unaffected by light treatment. The results are discussed in terms of the differences in function of vertically and horizontally oriented spacers, and the probable benefits of plasticity in enhancing essential activities including photosynthesis, habitat exploration and seed dispersal. Received: 23 December 1996 / Accepted: 19 July 1997     

Differential predation by Coleomegilla maculata on Colorado potato beetle strains that vary in growth on tomato

This study tests the hypothesis that the generalist predator Coleomegilla maculata DeGeer causes differential mortality of Colorado potato beetle, Leptinotarsa decemlineata (Say), larvae differing in their degree of genetic adaptation to tomato (Lycopersicon esculentum Mill.) as a host plant. Results of a series of laboratory experiments demonstrate that adult C. maculata can cause higher mortality to nonadapted than adapted Colorado potato beetle larvae. The extent of differential mortality caused by C. maculata depended on age of potato beetle larvae; presence of potato beetle eggs; whether or not the predator had a choice among prey items; and, in choice situations, the ratio of adapted to nonadapted potato beetle larvae. Although adult C. maculata have the potential to prey differentially on tomato-adapted and nonadapted Colorado potato beetle larvae in mixed populations, the magnitude of differential predation in a natural setting could be highly variable.     

Shade avoidance in the clonal herb Trifolium fragiferum/: a field study with experimentally manipulated vegetation height

In herbaceous canopies light availability can show high degrees of spatial variability in a vertical and also in a horizontal direction. Stoloniferous plants are hence likely to encounter differences in light availability during their ontogenetic development. Different mechanisms, such as petiole elongation, plasticity in internode length and branching, and an enhanced allocation to sexual reproduction have been suggested to represent viable shade-avoidance mechanisms for clonal plants.In a field experiment we tested the response of the stoloniferous herb Trifolium fragiferum L. to experimentally manipulated vegetation heights. Naturally occurring clonal fragments were exposed to four different vegetation heights ranging from 0 cm (high light availability created by clipping the surrounding natural vegetation at ground level) to 20 cm (natural shading in closed canopy). The growth and development of individual clones was followed for two months. At the end of the experiment above-ground plant parts were harvested. Growth-related and morphological parameters (e.g., petiole and internode length) as well as patterns of meristem utilization (i.e., flowering, branching) were recorded.Neither primary stolon growth and biomass accumulation nor branching and flowering were significantly affected by treatments. However, increased vegetation height resulted in a reduced number of secondary ramets and also had strong positive effects on petiole length, leading to marked changes in the architecture of plants growing in canopies of different heights. In addition, the average weight of individual ramets on the primary stolon was markedly higher in plants exposed to taller vegetation as compared to shorter vegetation.The results of this study suggest the occurrence of a trade-off between clonal expansion (i.e., secondary ramet production) and the average size of clonal offspring. If grown under higher vegetation plants invested more into the size of individual ramets, especially into elongating petioles, and less into the growth and development of lateral branches. Placing leaf laminae higher up in the canopy results in an enhanced light interception which has apparently buffered negative effects of increased vegetation height on whole-clone biomass. Plants grown under shorter vegetation invested more into lateral spread by producing more, but smaller ramets.     

Effects of cadmium on integration and resource allocation in the clonal fern Salvinia molesta

Summary In clonal plants exposed to pollution, ramets which are physiologically integrated may be less fit than ramets which are independent, if (a) translocation of toxins from contaminated ramets produced toxicity, or (b) toxicity in parent ramets reduced the degree of nutritional support to otherwise non-exposed daughters. These hypotheses were tested in the aquatic fern Salvinia molesta exposed to cadmium. Pre-treatment of parent ramets with cadmium decreased the number and biomass of daughters subsequently produced in a cadmiumfree medium, because of reduced parental support of the first daughter generation. Second generation and later daughters were unaffected. Pre-treatment did not affect the pattern of integration (which, in terms of apical daughters' biomass, was bimodal with increasing colony size), or concentrations of essential elements in new growth. However, a diversion of resources from lateral to apical daughters occurred as a result of pre-treatment, especially in colonies with 3 attached parents. Loss/gain analysis showed that the diversion was almost reciprocal in terms of biomass, ramet numbers and phosphorus content. Integration between contaminated and uncontaminated ramets was not disadvantageous to the clone as a whole. However, integration was disadvantageous for Ca, Mg and Zn concentrations in daughters, which declined 15–22%. Because of enhanced apical growth, an indirect benefit of integration may be a more rapid fragmentation and dispersal of daughters from the site of contamination than if the parents were independent.     

Stochastic simulation of clonal growth in the tall goldenrod,Solidago altissima

Summary As clonal plants grow they move through space. The movement patterns that result can be complex and difficult to interpret without the aid of models. We developed a stochastic simulation model of clonal growth in the tall goldenrod, Solidago altissima. Our model was calibrated with field data on the clonal expansion of both seedlings and established clones, and model assumptions were verified by statistical analyses.When simulations were based on empirical distributions with long rhizome lengths, there was greater dispersal, less leaf overlap, and less spatial aggregation than when simulations were based on distributions with comparatively short rhizome lengths. For the field data that we utilized, variation in rhizome lengths had a greater effect than variation for either branching angles or rhizome initiation points (see text). We also found that observed patterns of clonal growth in S. altissima did not cause the formation of fairy rings. However, simulations with an artificial distribution of branching angles demonstrate that fairy rings can result solely from a plant's clonal morphology.Stochastic simulation models that incorporated variation in rhizome lengths, branching angles, and rhizome initiation points produced greater dispersal and less leaf overlap than deterministic models. Thus, variation for clonal growth parameters may increase the efficiency of substrate exploration by increasing the area covered and by decreasing the potential for intraclonal competition. We also demonstrated that ramet displacements were slightly, but consistently lower in stochastic simulation models than in random-walk models. This difference was due to the incorporation of details on rhizome bud initiation into stochastic simulation models, but not random-walk models. We discuss the advantages and disadvantages of deterministic, stochastic simulation, and random-walk models of clonal growth.     

Effects of light quality and quantity on growth of the clonal plant Eichhornia crassipes

Summary Plant canopy shade reduces photosynthetic photon flux density (PPFD) and ratio of red to far-red light (z). Both effects can cause plants to increase potential for light acquisition through vertical growth and leaf area expansion. Clonal plants such as Eichhornia crassipes might alternatively increase light interception via horizontal growth of stolons or rhizomes and placement of new ramets in less shaded microsites. Effect of simulated canopy shade and component effects of PPFD and z were tested by filtering or adding light uniformly, to a whole group of connected ramets, or locally, to individual ramets within a group. In uniform treatments, low PPFD reduced total growth but low z did not. Low PPFD and low z independently reduced stolon and ramet production and caused etiolation of petioles; effect of low PPFD plus low z on ramet production was greater than that of either factor alone. Lateral clonal growth thus did not seem to be a response to uniform shading; instead, uniformly low PPFD or low z increased partitioning to established ramets. Low z changed partitioning without changing total growth. In local treatments, reduction of growth of individual ramets due to low PPFD and inhibition of new ramet production attributable to spectral composition of light were mitigated when connected ramets were unshaded; plants may respond differently to patchy than to uniform shade.     

Effects of water depth and substrate type on growth dynamics and biomass allocation of Oryza rufipogon

Oryzarufipogon is known as the ancestor of Asian cultivated rice (Oryzasativa) and the most important germplasm for rice improvement. In this study, we investigated its growth and biomass allocation in response to water depth and substrate type. The results demonstrated that individual plant had an average weight of 18.98 g, 9.36 g, and 10.59 g and produced 77.7 seeds, 35.5 seeds, and 41.6 seeds in deep-, fluctuated-, and shallow water, respectively. Root-to-shoot ratios were 0.37 in deep water, 0.61 in fluctuated water level, and 0.56 in shallow water, reflecting larger investment in roots under water-level fluctuation and shallow water conditions. However, biomass allocation for sexual reproduction was no significant different among the three water levels. In substrates experiment, total biomass and seed production were significantly different between sediment types, and the averages were 10.59 g, 3.56 g, and 0.76 g in biomass and 41.6 seeds, 13.8 seeds, and 0.5 seeds per plant in lake sediment, river sand and clay, respectively. The number of tillers was significantly fewer in sandy and clay substrates than in lake sediments. Individual plant in clay substrates was significantly shorter than those in sandy and in lake sediment substrates. Our study suggests that the plasticity in morphology and biomass allocation of O.rufipogon appears to be an adaptation to the variability in water level and substrate conditions.     

Effects of simulated root herbivory and fertilizer application on growth and biomass allocation in the clonal perennialSolidago canadensis

Summary Compensatory growth in response to simulated belowground herbivory was studied in the old-field clonal perennialSolidago canadensis. We grew rootpruned plants and plants with intact root systems in soil with or without fertilizer. For individual current shoots (aerial shoot with rhizome and roots) and for whole clones the following predictions were tested: a) root removal is compensated by increased root growth, b) fertilizer application leads to increased allocation to aboveground plant organs and increased leaf turnover, c) effects of fertilizer application are reduced in rootpruned plants. When most roots (90%) were removed current shoots quickly restored equilibrium between above-and belowground parts by compensatory belowground growth whereas the whole clone responded with reduced aboveground growth. This suggests that parts of a clone which are shared by actively growing shoots act as a buffer that can be used as source of material for compensatory growth in response to herbivory. Current shoots increased aboveground mass and whole clones reduced belowground mass in response to fertilizer application, both leading to increased allocation to aboverground parts. Also with fertilizer application both root-pruned and not root-pruned plants increased leaf and shoot turnover. Unfertilized plants, whether rootpruned or not, showed practically no aboveground growth and very little leaf and shoot turnover. Effects of root removal were as severe or more severe under conditions of high as under conditions of low nutrients, suggesting that negative effects of belowground herbivory are not ameliorated by abundant nutrients. Root removal may negate some effects of fertilizer application on the growth of current shoots and whole clones.     

Effects of water depth and substrate type on growth dynamics and biomass allocation of Oryza rufipogon

Oryzarufipogon is known as the ancestor of Asian cultivated rice (Oryzasativa) and the most important germplasm for rice improvement. In this study, we investigated its growth and biomass allocation in response to water depth and substrate type. The results demonstrated that individual plant had an average weight of 18.98 g, 9.36 g, and 10.59 g and produced 77.7 seeds, 35.5 seeds, and 41.6 seeds in deep-, fluctuated-, and shallow water, respectively. Root-to-shoot ratios were 0.37 in deep water, 0.61 in fluctuated water level, and 0.56 in shallow water, reflecting larger investment in roots under water-level fluctuation and shallow water conditions. However, biomass allocation for sexual reproduction was no significant different among the three water levels. In substrates experiment, total biomass and seed production were significantly different between sediment types, and the averages were 10.59 g, 3.56 g, and 0.76 g in biomass and 41.6 seeds, 13.8 seeds, and 0.5 seeds per plant in lake sediment, river sand and clay, respectively. The number of tillers was significantly fewer in sandy and clay substrates than in lake sediments. Individual plant in clay substrates was significantly shorter than those in sandy and in lake sediment substrates. Our study suggests that the plasticity in morphology and biomass allocation of O.rufipogon appears to be an adaptation to the variability in water level and substrate conditions.     

The effect of local resource availability and clonal integration on ramet functional morphology in Hydrocotyle bonariensis

Summary Within a physiologically integrated clone, the structure and functioning of an individual ramet is determined by: 1) the response of that ramet to its local environment and 2) its response to resource integration within the clone. In a multifactorial experiment, Hydrocotyle bonariensis ramets were grown in limiting resource environments with and without the benefit of basipetal resource movement from another branch of the clone. Ramets were analyzed for their morphological responses to variation in local light, water and nitrogen availability and to the superimposed effect of resource integration on these conditions. The expression of ramet morphology, from induction to development, was highly plastic in response to variable local resource availability. Resource integration changed a ramet's local response in a variety of ways depending on the resource(s) being translocated and the character involved. Among leaf characteristics (leaf weight, petiole height, blade area), resource translocation into the shade resulted in an enhancement of the local response. Similarly, the translocation of nitrogen and water generally increased clonal proliferation and sexual reproduction among ramets. In contrast, the translocation of water reversed the effect of local low water conditions on ramets by inhibiting root production. Some characters such as internode distance and leaf allometry were unaffected by integration. The maintenance of connections between ramets as a Hydrocotyle clone expands allows for resource sharing among widely separated ramets and can result in an integrated morpological response to a resource environment that is patchy in time and space.     

不同光环境对海南龙血树幼苗表型可塑性及生存策略的影响

为了解光照对海南龙血树(Dracaena cambodiana)幼苗生长的影响,研究了不同光照环境下海南龙血树幼苗形态、生理特性和生物量分配的变化,并分析了其生态适应性。结果表明,海南龙血树幼苗的形态、生理和生物量分配指标在不同光照强度间存在显著差异,各指标的可塑性指数为0.08~0.86,其中根茎叶及总生物量的可塑性指数普遍较高(0.67~0.86),表明海南龙血树幼苗有较好的光照适应性,其策略主要是通过调整根茎叶生物量的分配来适应光照的变化。随着光照强度的降低,海南龙血树幼苗的比叶面积、叶根比呈现显著增大趋势,表明幼苗可通过增加单株叶面积比例,扩大光合作用面积,有效调节自身生物量配置。37.3%自然光照(L2)是海南龙血树幼苗生长的最佳光照强度。现存海南龙血树生境改变,生境缺少林荫以致光照强度过大,不利于幼苗根系生长,难以度过干旱季节,可能是海南龙血树自然更新失败的重要原因之一。     

Competitive ability of two Brassica varieties in relation to biomass allocation and morphological plasticity under varying nutrient availability

Green cabbage (Brassica campestris, leafy variety) and turnip (Brassica campestris var. rapifera, rooty variety) were grown in both monocultures and mixtures at three nutrient levels to investigate their responses to nutrient availability with respect to biomass allocation, morphological plasticity and competitive ability. Their allocation parameters and leaf morphological traits were affected by both nutrient availability and developmental stage. Both of the varieties had a smaller biomass allocation to leaf blades, but a greater allocation to petioles at high nutrient levels. Root:shoot ratio (RSR) of green cabbage decreased with increasing nutrient availability, whereas that of turnip increased. Turnip had a smaller leaf blade weight ratio (LBWR) than cabbage, being compensated for by larger leaf area ratio (LAR) and specific leaf area (SLA). Leaf area ratio and SLA of both the varieties increased with increasing nutrient availability as did their mean dry weights. The mean dry weight of turnip was slightly greater than that of green cabbage in their respective monocultures, while that of green cabbage was greater than that of turnip in their 1:1 mixture. Therefore, green cabbage, having inherently greater biomass allocation to leaves, was generally more competitive than turnip with more biomass allocation to roots, especially at higher nutrient levels. However, within a variety, morphological plasticity (variation in LAR and SLA) was more important than the plasticity in biomass allocation (e.g. variation in RSR and LBWR) in determining competitive ability. The implication of our results is that competition models based on biomass allocation pattern alone may fail to predict competitive outcomes and that such models should also take morphological plasticity into full account.     

光照对苦瓜形态可塑性及生物量配置的影响

在人为遮阴条件下,对苦瓜的生长动态、形态特征以及生物量分配进行了研究。结果表明,不同遮阴处理下苦瓜植株的生长有较大的差异,弱光照不利于苦瓜构件数量的增加和生物量的积累;植株在弱光下形成较少的分枝,较薄的叶片以及较细长的主茎和叶柄,表现出较强光生长环境下更强的形态可塑性;植株在生长早期较生长晚期有较大的形态可塑性;生物量对叶片和主茎的分配随光照的减弱而增大,分枝的生物量分配随光照的减弱而降低,光照对分枝茎的生物量分配影响不大;在光资源充足的情况下,外界支持物的缺乏不会对苦瓜的生长造成太大的影响。     

Multiple forms of phycoerythrin-545 from Cryptomonas maculata

Three multiple phycoerythrin-545 forms were purified from crude extracts of Cryptomonas maculata by preparative isoelectric focusing. The phycoerythrin forms are charge isomers with isoelectric points at 7.83, 5.05 and 4.84. The multiple pigment forms have similar molecular weights of 44500 daltons and are composed of subunits of unequal size in a 1:1 stoichiometry with molecular weights of () 9900 and () 15700 daltons, twice. The proposed quarternary structure of the native pigments is ()₂()₂.The charge differences of the phycoerthrins are caused by a charge heterogeneity of the light subunits, as revealed by urea gel electrophoresis. The chains of pigment form pI 7.83 had a greater electrophoretic mobility than those subunits of the acidic pigment forms pI 5.05 and pI 4.84.The phycoerythrin forms have an absorption spectrum with similar absorption maxima at 544 nm, but differ in the position of the long wavelength shoulders lying at 555 and 557 nm in the negatively charged pigment forms and at 560 nm for the phycoerythrin form with a pI at 7.83.The fluorescence emission spectra coincide in their asymmetrical shape with shoulders at about 620 nm; they slightly differ int he position of the emission maxima at 586 nm for the phycoerythrins with pIs at 4.84 and 5.05 and at 584 nm for phycoerythrin with pI at 7.83.Abbreviations PC phycocyanin - PE phycoerythrin - pI isoelectric point - SDS sodium dodecyl sulphate     

Potential and limitations of current concepts regarding the response of clonal plants to environmental heterogeneity

Plant ecologists have spent considerable effort investigating the physiological mechanisms and ecological consequences of clonal growth in plants. One line of research is concerned with the response of clonal plants to environmental heterogeneity. Several concepts and hypotheses have been formulated so far, suggesting that intra-clonal resource translocation, morphological plasticity on different organizational levels (e.g. leaves, ramets, fragments), and other features of clonal plants may represent potentially adaptive traits enabling stoloniferous and rhizomatous species to cope better with habitat patchiness. Although each of these concepts contributes substantially to our understanding of the ecology of clonal species, it is difficult to combine them into a consistent theoretical framework. This apparent lack of conceptual coherence seems partly be caused by an uncritical use of the term habitat heterogeneity. Researchers have not always acknowledged the fact that heterogeneity may refer to a number of fundamentally different aspects of environmental variability (i.e. scale, contrast, predictability, temporal vs. spatial heterogeneity), and that each of these aspects may, on one hand, allow for the evolution of specific plant responses to heterogeneity and, on the other, severely constrain the viability of potentially adaptive traits. Since adaptive responses are operational only in a narrow range of conditions (delimited by external environmental conditions and constraints internal to plants) it seems imperative to clearly define the context and the limits within which concepts regarding clonal plants' responses to heterogeneity are valid. In this paper an attempt is made to review a number of these concepts and to try and identify the necessary conditions for them to be operational. Special attention is paid (1) to different aspects of environmental heterogeneity and how they may affect clonal plants, and (2) to possible constraints (e.g. sectoriality, perception of environmental signals, morphological plasticity) on plant responses to patchiness.