Interspecific Variation in RGR and the Underlying Traits among 24 Grass Species Grown in Full Daylight

Abstract: A growth analysis was conducted with 24 central European grass species in full daylight to test whether traits underlying interspecific variation in relative growth rate (RGR) are the same in full daylight as they are at lower light, and whether this depends on the ecological characteristics of the studied species, i.e., their requirements with respect to nutrient and light availability.
In contrast to studies with herbaceous species at lower light, net assimilation rate (NAR) contributed more than leaf area ratio (LAR) or specific leaf area (SLA) to interspecific variation in RGR. This was associated with a larger interspecific variation in NAR than found in experiments with lower light. Without the two most shade-tolerant species, however, the contribution of LAR and its components to interspecific variation in RGR was similar or even higher than that of NAR.
Leaf dry matter content correlated negatively with RGR and was the only component of LAR contributing in a similar manner to variation in LAR and RGR. There was a positive correlation between NAR and biomass allocation to roots, which may be a result of nutrient-limited growth. RGR correlated negatively with biomass allocation to leaves. Leaf thickness did not correlate with RGR, as the positive effect of thin leaves was counterbalanced by their lower NAR.
Low inherent RGR was associated with species from nutrient-poor or shady habitats. Different components constrained growth for these two groups of species, those from nutrient-poor habitats having high leaf dry matter content, while those from shady habitats had thin leaves with low NAR.     

Differences in seedling growth behaviour among species: trait correlations across species, and trait shifts along nutrient compared to rainfall gradients

1 Species-pairs from woody dicot lineages were chosen as phylogenetically independent contrasts (PICs) to represent evolutionary divergences along gradients of rainfall and nutrient stress, and within particular habitat types, in New South Wales, Australia. Seedlings were grown under controlled, favourable conditions and measurements were made for various growth, morphological and allocation traits.
2 Trait correlations across all species were identified, particularly with respect to seedling relative growth rate (RGR) and specific leaf area (SLA), a fundamental measure of allocation strategy that reflects the light-capture area deployed per unit of photosynthate invested in leaves.
3 Across all species, SLA, specific root length (SRL) and seed reserve mass were the strongest predictors of seedling RGR. That is, a syndrome of leaf and root surface maximization and low seed mass was typical of high RGR plants. This may be a high-risk strategy for individual seedlings, but one presumably mitigated by a larger number of seedlings being produced, increasing the chance that at least one will find itself in a favourable situation.
4 Syndromes of repeated attribute divergence were identified in the two sets of gradient PICs. Species from lower resource habitats generally had lower SLA. Thus, in this important respect the two gradients appeared to be variants of a more general 'stress' gradient.
5 However, trends in biomass allocation, tissue density, root morphology and seed reserve mass differed between gradients. While SLA and RGR tended to shift together along gradients and in within-habitat PICs, no single attribute emerged as the common, primary factor driving RGR divergences within contrasts. Within-habitat attribute shifts were of similar magnitude to those along gradients.     

Measuring the RGR of Individual Grass Plants

Vegetative growth of grasses was analysed by dry mass increaseof growing leaves.Holcus lanatuswas grown in a controlled environmentand leaf extension rates of leaf numbers 5–10 of the maintiller were monitored daily. Leaf appearance and leaf extensionrates (LER) of leaves 5–7 enabled the prediction of thefinal length and dry mass of leaf 8 during its growth. A linearincrease of leaf mass per unit leaf length (LML) of leaf 8 wasobserved during growth. After harvest the daily increase indry mass of growing leaves was calculated from the LER and correspondingincrease in LML. The relative growth rate (RGR) of the maintiller showed day-to-day fluctuations and was gradually reducedby 50% over a 16-d period. The RGR of the shoot was maintainedby tillering. The RGR of a single (grass) plant can be calculatedfrom four parameters only: LER, LML, leaf appearance and tillering.Variation of RGR over a period can be reconstructed after harvestand the impact of these four parameters on RGR can be established.Copyright1998 Annals of Botany Company. Relative growth rate, grass, leaf growth,Holcus lanatus.     

Phenolic and phenolic-related factors as determinants of suitability of mountain birch leaves to an herbivorous insect

We investigated the role of phenolic and phenolic-related traits of the leaves of mountain birch (Betula pubescens ssp. czerepanovii) as determinants of their suitability for the growth of larvae of the geometrid Epirrita autumnata. As parameters of leaf suitability, we determined the contents of total phenolics, gallotannins, soluble and cell-wall-bound proanthocyanidins (PAS and PAB, respectively), lignin, protein precipitation capacity of tannins (PPC), and leaf toughness. In addition, we examined concentrations of soluble carbohydrates and protein-bound amino acids as background variables describing the nutritive value of leaves. The correlation of the leaf traits of our 40 study trees with the tree-specific relative growth rate (RGR) of E. autumnata showed that the only significant correlation with RGR was that of PAS - the largest fraction of total phenolics - and even that explained only 15% of the variation in E. autumnata growth. The nonlinear estimation of the relationship between RGR and PAS by piecewise linear regression divided the 40 study trees into two groups: (i) 19 trees with good leaves for E. autumnata (RGR ranging from 0.301 to 0.390), and (ii) 21 trees with poor leaves (RGR ranging from 0.196 to 0.296). The suitability of leaves within these two groups of trees was determined by different phenolic traits. Within the good group, the suitability of leaves for larvae was determined by the PPC of extracts, which strongly correlated with gallotannins, and by the total content of gallotannins. In contrast, the leaves of poor trees had significantly higher contents of both PAS and PAB, but leaf toughness correlated only negatively with the RGR of E. autumnata larvae. We also discuss the causes of variation in the phenolic and phenolic-related factors that determine the suitability of leaves for E. autumnata larvae in different groups of trees.     

岷江上游干旱河谷矮探春叶片特征与环境因子的关系

对岷江上游干旱河谷矮探春(Jasminumhumile L.)的叶片形态解剖特征进行了显微观察,分析了海拔梯度上叶片形态与环境因子的关系。结果表明,矮探春叶片呈长椭圆形,叶肉组织分化明显;随着海拔升高,叶面积、厚度、干重、饱和含水量、海绵组织厚度,中脉厚度以及厚角组织厚度/中脉厚度之比(M/C)等呈增大趋势,而栅栏组织厚度/海绵组织厚度之比(P/S)则减小;叶片长/宽比、表皮厚度、栅栏组织厚度和比叶重在海拔梯度上无明显差异。叶面积、干重、饱和含水量、叶厚度和海绵组织厚度等参数两两之间呈显著正相关,而它们与P/S均呈显著负相关。叶面积、干重、海绵组织厚度和M/C等主要受土壤含水量的影响,并随着土壤含水量增加而增大;P/S随着土壤含水量和年降水量增加而减小;叶片厚度、饱和含水量和中脉厚度均随着温和度的降低而增大。岷江上游干旱河谷区土壤水份和生长季温度可能是影响矮探春叶片形态解剖特征的主要因子。     

Scaling sun and shade photosynthetic acclimation of Alocasia macrorrhiza to whole-plant performance – II. Simulation of carbon balance and growth at different photon flux densities

A whole-plant carbon balance model incorporating a light acclimation response was developed for Alocasia macrorrhiza based on empirical data and the current understanding of light acclimation in this species. The model was used to predict the relative growth rate (RGR) for plants that acclimated to photon flux density (PFD) by changing their leaf type, and for plants that produced only sun or shade leaves regardless of PFD. The predicted RGR was substantially higher for plants with shade leaves than for those with sun leaves at low PFD. However, the predicted RGR was not higher, and in fact was slightly lower, for plants with sun leaves than for those with shade leaves at high PFD. The decreased leaf area ratios (LARs) of the plants with sun leaves counteracted their higher photosynthetic capacities per unit leaf area (A_max). The model was manipulated by changing parameters to examine the sensitivity of RGR to variation in single factors. Overall, RGR was most sensitive to LAR and showed relatively little sensitivity to variation in A_max or maintenance respiration. Similarly, RGR was relatively insensitive to increases in leaf life-span beyond those observed. Respiration affected RGR only at low PFD, whereas A_max was moderately important only at high PFD.     

罗汉松雌雄株叶形态结构的比较研究

该研究以雌雄异株植物罗汉松(Podocarpus macrophyllus)成熟叶为研究材料,采用光学显微镜、扫描电镜和透射电镜观察比较罗汉松雌、雄植株叶在形态、显微结构和超显微结构的差异,以明确罗汉松雌、雄株在进化过程中叶对环境功能的适应性。结果显示:(1)罗汉松雌株的叶片大于雄株,且两者的叶长、叶宽和叶柄长差异极显著,而叶柄厚、叶面积、叶体积、叶质量、比叶重(SLW)、面积与体积之比(A/V)等性状无显著差异。(2)雌株叶片的气孔相对较大,密度较高,且雌株气孔宽度极显著大于雄株;雌株叶片的上表皮长细胞宽度和下表皮短细胞宽度均显著大于雄株,但雌株叶片的上表皮长细胞和短细胞的长度则显著小于雄株。(3)罗汉松雌株叶片的栅栏组织厚度、海绵组织厚度、传输组织长度和宽度、上下角质层厚度、维管束厚度、叶片紧密度(CTR)及疏松度(SR)均极显著大于雄株,而雌株的下表皮厚度极显著小于雄株,但雌雄株叶片的上表皮细胞厚度和栅海比差异不显著;雌株叶片的栅栏组织细胞、叶绿体和线粒体均较雄株的长而细,且雌株的线粒体宽度极显著小于雄株。(4)罗汉松雌株叶片上表皮蜡质饰纹、下表皮角质层纹饰、气孔外拱盖纹饰及内缘类型等4个微形态特征与雄株差异明显。(5)叶表皮蜡质层能谱分析表明,罗汉松雌株叶片含有9种元素,而雄株叶片仅有8种(缺少K元素);且雌株的Si元素含量高于雄株,而雄株的C、O、Na、Mg、Al、Ca和Au元素含量均高于雌株。研究表明,罗汉松雌、雄植株之间存在明显的第二性征,雌株叶片结构有助于提高光合等性能以满足生殖需求;罗汉松雌、雄株叶形态结构的差异是其长期进化形成的有利于物种繁衍的适应策略。     

Latitudinal variation in plant size and relative growth rate in Arabidopsis thaliana

Latitude is an important determinant of local environmental conditions that affect plant growth. Forty ecotypes of Arabidopsis thaliana were selected from a wide range of latitudes (from 16°N to 63°N) to investigate genetic variation in plant size and relative growth rate (RGR) along a latitudinal gradient. Plants were grown in a greenhouse for 31 days, during which period three consecutive harvests were performed. Plants from high latitudes tended to have smaller plant size in terms of seed size, cotyledon width, rosette size, number of rosette leaves, size (leaf area) of the largest leaves, total leaf area, and total dry weight per plant than those from low latitudes. The mean (±SE) RGR across ecotypes was 0.229 (±0.0013) day⁻¹. There was, however, significant ecotypic variation, with RGR being negatively correlated with latitude. The two main components of RGR, leaf area ratio (LAR) and unit leaf rate (ULR), were also correlated with latitude: LAR increased with increasing latitude while ULR decreased with increasing latitude. It was also found that RGR tended to be negatively correlated with LAR, specific leaf area (SLA) and specific root length (SRL) but to be positively correlated with mean area per leaf (MAL) and ULR. The variation in RGR among ecotypes was relatively small compared with that in the other traits. RGR may be a conservative trait, whose variation is constrained by the trade-off between its physiological (i.e. ULR) and morphological (i.e. LAR) components. Received: 2 November 1997 / Accepted: 28 February 1998     

Relative growth rate variation of evergreen and deciduous savanna tree species is driven by different traits

Background and Aims

Plant relative growth rate (RGR) depends on biomass allocation to leaves (leaf mass fraction, LMF), efficient construction of leaf surface area (specific leaf area, SLA) and biomass growth per unit leaf area (net assimilation rate, NAR). Functional groups of species may differ in any of these traits, potentially resulting in (1) differences in mean RGR of groups, and (2) differences in the traits driving RGR variation within each group. We tested these predictions by comparing deciduous and evergreen savanna trees.

Methods

RGR, changes to biomass allocation and leaf morphology, and root non-structural carbohydrate reserves were evaluated for juveniles of 51 savanna species (34 deciduous, 17 evergreen) grown in a common garden experiment. It was anticipated that drivers of RGR would differ between leaf habit groups because deciduous species have to allocate carbohydrates to storage in roots to be able to flush leaves again, which directly compromises their LMF, whereas evergreen species are not subject to this constraint.

Key Results

Evergreen species had greater LMF and RGR than deciduous species. Among deciduous species LMF explained 27 % of RGR variation (SLA 34 % and NAR 29 %), whereas among evergreen species LMF explained between 2 and 17 % of RGR variation (SLA 32–35 % and NAR 38–62 %). RGR and LMF were (negatively) related to carbohydrate storage only among deciduous species.

Conclusions

Trade-offs between investment in carbohydrate reserves and growth occurred only among deciduous species, leading to differences in relative contribution made by the underlying components of RGR between the leaf habit groups. The results suggest that differences in drivers of RGR occur among savanna species because these have different selected strategies for coping with fire disturbance in savannas. It is expected that variation in the drivers of RGR will be found in other functional types that respond differently to particular disturbances.     

Toward a causal explanation of plant invasiveness: seedling growth and life-history strategies of 29 pine (Pinus) species

We studied 29 pine (Pinus) species to test the hypothesis that invasive species in disturbed habitats have distinct attributes. Seedling relative growth rate (RGR) and measures of invasiveness were positively associated across species as well as within phylogenetically independent contrasts. High RGR, small seed masses, and short generation times characterize pine species that are successful invaders in disturbed habitats. Discriminant analysis and logistic regression revealed that RGR was the most significant factor among these life-history traits separating invasive and noninvasive species. We also explored the causes of differences in RGR among invasive and noninvasive species. While net assimilation rate, leaf mass ratio, and specific leaf area (SLA) were all found to be contributing positively to RGR, SLA was found to be the main component responsible for differences in RGR between invasive and noninvasive pines. We investigated differences in SLA further by studying leaf anatomy, leaf density, and leaf thickness. We also evaluated relative leaf production rate as an important aspect of SLA. We proposed a hypothetical causal network of all relevant variables.     

Morphological,structural and physiological differences in heteromorphic leaves of Euphrates poplar during development stages and at crown scales

• Euphrates poplar (Populus euphratica Oliv.) has heteromorphic leaves including strip, lanceolate, ovate, and broad‐ovate leaves from base to top in the mature canopy.
• To clarify how diameter at breast height (DBH) and tree height affect the functional characteristics of all kinds of heteromorphic leaves, we measured the morphological anatomical structure and physiological indices of five crown heteromorphic leaves of P. euphratica at 2, 4, 6, 8, 10, and 12 m from the same site. We also analysed the relationships between morphological structures and physiological characteristics of heteromorphic leaves and DBH and the height of heteromorphic leaves.
• The results showed that the number of abnormalities regarding blade width, leaf area, leaf thickness, leaf mass per area, cuticle layer thickness, palisade tissue thickness, and palisade tissue/sponge tissue ratio increased with size order and sampling height gradient. Net photosynthetic rate, transpiration rate, stomatal conductance, instantaneous water use efficiency, stable delta carbon isotope ratio, proline and malondialdehyde (MDA) content increased with DBH and sampling height. By contrast, blade length, leaf shape index, and intercellular CO2 concentration decreased with the increase in path order and sampling height gradient. Although MDA content and leaf sponge thickness were not correlated with DBH or sampling height, other morphological structure and physiological parameters were significantly correlated with these variables. In addition, correlations were found among leaf morphology, anatomical structure, and physiological index parameters indicating that they changed with path order and tree height gradient.
• The differences in the morphology, anatomic structure and physiological characteristics of the heteromorphic leaves ofP. euphratica are related to ontogenesis stage and coronal position.

     

Variation in relative growth rate and its components in the annual Polygonum aviculare in relation to habitat disturbance and seed size

Polygonum aviculare is an annual weedy species showing extensive genetic variation in seed and leaf size and colonizing various types of man-disturbed habitats. A growth analysis was conducted on 12 genotypes representative of three regimes of disturbance of natural habitat (trampling, weeding, and no disturbance in the course of the growing season), grown under productive conditions in order to test whether relative growth rate (RGR) varies at the intraspecific level and, if so, which growth parameters may explain its variation. RGR showed significant genotypic variation (0.355–0.452 g g^-1 day^-1), positively correlated with specific leaf area (SLA) and leaf mass ratio (LMR) and negatively correlated with unit leaf rate per unit leaf area (ULR_A). Thus, the paramount importance of leaf area ratio (LAR=SLA×LMR) in determining growth rate variation between different herbaceous species is confirmed at the intraspecific level in this species. Genotypes originating from trampled habitats had smaller seeds and smaller leaves than genotypes from habitats subject to other disturbance regimes. Additionally, they showed a lower LAR, not entirely compensated for by a higher ULR_A, which resulted in a positive allometric relationship between seed size and RGR. It is hypothesized that their lower SLA, correlated with a higher leaf dry matter content (possibly a consequence of a higher cell wall content per unit leaf area) and their lower LMR have been co-selected with small leaf size as adaptations promoting resistance to trampling stress. It is suggested that variation in cell size and/or gibberellin content might be the mediators of the correlation found between seed size, leaf size and growth parameters within this species.     

Intra-specific variation in relative growth rate: impact on competitive ability and performance of Lychnis flos-cuculi in habitats differing in soil fertility

Plant species from unproductive or adverse habitats are often characterized by a low potential relative growth rate (RGR). Although it is generally assumed that this is the result of selection for specific trait combinations that are associated with a low rate of net biomass accumulation, few studies have directly investigated the selective (dis-)advantage of specific growth parameters under a set of different environmental conditions. Aim of the present study was to quantify the impact of inherent differences in growth parameters among phenotypes of a single plant species, Lychnis flos-cuculi, on their performance under different soil nutrient conditions. Growth analysis revealed significant variation in RGR among progeny families from a diallel cross between eight genotypes originating from a single population. Differences in RGR were due to variation in both leaf area ratio (LAR) and in net assimilation rate (NAR). A genetic trade-off was observed between these two components of growth, i.e. progeny families with high investment in leaf area had a lower rate of net biomass accumulation per unit leaf area. The degree of plasticity in RGR to nutrient conditions did not differ among progeny families. Inherent differences in growth parameters among progeny families had a significant impact on their yield in competition with Anthoxanthum odoratum and Taraxacum hollandicum. In nutrient-rich conditions, progeny families with an inherently high leaf weight ratio (LWR) achieved higher yield in competition, but variation in this trait could not explain differences in competitive yield under nutrient-poor conditions. Inherent differences in growth parameters among progeny families were poorly correlated with differences in survival and average rosette biomass (a good predictor of fecundity) among these progeny families sown in four field sites along a natural gradient of soil fertility. In the more productive sites none of the growth parameters was significantly correlated with rosette biomass, but in the least productive site progeny families with an inherently high specific leaf area (SLA) tended to produce smaller rosettes than low-SLA families. These results are consistent with the view that a selective advantage may accrue from either high or low values of individual RGR components, depending on habitat conditions, and that the selective advantage of low trait values in nutrient-poor environments may results in indirect selection for low RGR in these habitats.     

Traits of an invasive grass conferring an early growth advantage over native grasses

Aims Invasive species often have higher relative growth rates (RGR) than their native counterparts. Nutrient use efficiency, total leaf area and specific leaf area (SLA) are traits that may confer RGR differences between natives and invasives, but trait differences are less prominent when the invasive species belongs to the same plant functional type as the dominant native species. Here, we test if traits displayed soon after germination confer an early size advantage. Specifically, we predicted that invasive species seedlings grow faster than the natives because they lack trade-offs that more strongly constrain the growth of native species.Methods We quantified plant morphological and physiological traits and RGR during early seedling growth at high and low nutrient levels in three dominant perennial native C₄ grasses: Panicum virgatum L. (switchgrass), Schizachyrium scoparium (Michx.) Nash (little bluestem) and Andropogon gerardii Vitman (big bluestem); and a perennial C₄ exotic invasive grass, Sorghum halepense (L.) Pers. (Johnsongrass).Important findings After 2 weeks of growth, Johnsongrass seedlings had greater biomass, SLA and photosynthetic nitrogen use efficiency, but lower leaf N concentrations (% leaf N) and root:shoot ratio than natives. As growth continued, Johnsongrass more quickly produced larger and thicker leaves than the natives, which dampened the growth advantage past the first 2 to 3 weeks of growth. Investment in carbon gain appears to be the best explanation for the early growth advantage of Johnsongrass. In natives, growth was constrained by an apparent trade-off between allocation to root biomass, which reduced SLA, and production of leaves with high N content, which increased carbon gain. In Johnsongrass, root:shoot ratio did not interact with other traits, and % leaf N was decoupled from RGR as a result of a trade-off between the positive indirect association of % leaf N with RGR and the negative direct association of % leaf N with RGR.     

Factors controlling plasticity of leaf morphology in Robinia pseudoacacia: III. biophysical constraints on leaf expansion under long-term water stress

In this article, we measured the relative growth rate (RGR) of leaves of Robinia pseudoacacia seedlings under well-watered and water-stressed conditions (mid-day Ψ(w) = leaf water potential estimated with a pressure bomb of -0.48 and -0.98 MPa, respectively). Pressure-volume (PV) curves were done on growing leaves at 25, 50 and 95% of the mature size (growth stage) in order to compute solute potential (Ψ) and turgor pressure (Ψ(P) ) as a function of Ψ(w) . The PV curves and diurnal measurements of Ψ(w) and RGR allowed us to evaluate the parameters (cell wall extensibility m and growth turgor threshold Y) of the Lockhart equation, RGR = m(Ψ(P)-Y), at each growth stage. Our data showed that m and Y did change with leaf age, but the changes were slow enough to evaluate m and Y on any given day. We believe this is the first study to provide evidence that the Lockhart equation adequately quantifies leaf growth of trees over a range of time domains. The value of m linearly declined and Y linearly increased with growth stage. Also, mild drought stress caused a decline in m and increase in Y relative to controls. Although water stress caused an osmotic adjustment which, in turn, increased Ψ(P) in stressed plants relative to controls, the RGR and final leaf sizes were reduced in water-stressed plants because of the impact of water stress on decreased m and increased Y.     

Interaction of nitrogen and phosphorus nutrition in determining growth

In this paper we discuss the differences and similarities in the growth response of tomato plants to N and P limitation, and to their interaction. Two detailed growth experiments, with varied N or P supply, were conducted in order to unravel the effects of N and P limitation on growth of young tomato plants (Lycopersicon esculentum Mill.). Relative growth rate (RGR) initially increased sharply with increasing plant P concentration but leveled off at higher plant P concentrations. In contrast, RGR increased gradually with increasing plant N concentration before it leveled off at higher plant N concentrations. The relationship of RGR with organic leaf N and P showed the same shape as with total N and P concentrations, respectively. The difference in response is most likely due to the different roles of N and P in the machinery of the plant's energy metabolism (e.g., photosynthesis, respiration). Plant N concentration decreased with increasing P limitation. We show that this decrease cannot be explained by a shift in dry-mass partitioning. Our results suggest that the decrease in N concentration with increasing P limitation may be mediated by a decrease in leaf cytokinin levels and is less likely due to decreased energy availability at low P conditions. Dry-mass partitioning to the roots was closely linearly related to the leaf reduced-N concentration. However, treatments that were severely P limited deviated from this relationship.     

Control of Leaf Growth and its Role in Determining Variation in Plant Growth Rate from an Ecological Perspective

Abstract: Plants vary widely in their relative growth rate (RGR), be it dependent on environmental conditions or due to their genetic background. In a comparison of the RGR of grasses growing under different environmental conditions, variation in RGR tends to correlate with that in the leaf elongation rate (LER). When different species or genotypes thereof are compared under identical growing conditions, variation in LER may or may not correlate with that in RGR, depending on the comparison. However, since RGR is described by an exponential equation, whereas LER is mainly a linear process, we conclude that any correlation between RGR and LER must be fortuitous. That is, exponential growth must be due to increases with time in plant traits such as 1) leaf dry mass per unit leaf length invested per unit time, and/or 2), i.e., the total LER of all the growing leaves at one point in time. The latter can be achieved as follows: 1) each subsequent leaf has a higher LER than the preceding one; 2) leaves appear at an increasing rate; 3) the duration of the process of leaf elongation increases for subsequent leaves. In this review, we only explore possible factors that account for changes in with time, in different genotypes and under different environmental conditions. Inherent variation in LER of individual leaves and variation due to environmental factors may reflect variation in the rate of cell division and/or in cell elongation.     

Growth Analysis of Sumatran Monophyllaea, Possessing Only One Leaf Throughout Perennial Life

Abstract Growth and allometry were analyzed for populations of Monophyllaea hirtella Miq and M. horsfieldii R. Br. (Gesneriaceae), forest floor herbs that have only one cotyledonous leaf throughout life, in an equatorial rain forest in West Sumatra. Monophyllaea populations consisted of individuals of various sizes up to 30 g dry weight and 50x70 cm in leaf width and length. The relative growth rate (RGR) declined with size to an asymptotic value of 0.015–0.018 gg^-1 week^-1 for large individuals at sexual maturity (>2 g dry weight). The size-RGR relation did not differ among observations at three differen times of year and between two species in different habitals, indicating that it takes 4.6 years for seedlings to attain sexual maturity and 6.4 years to reach 10 g dry weight. Irrespective of embryonic organization of Monophyllaea , clear allometry existed among organs. Net assimilation rate was constant for juveniles and increased with size for adults. Decline of both the specific leaf area and the ratio of assimilate allocation to leaf caused the decrease of RGR with size. Reproductive allocation was 31% to reproductive organs and at most 5% to seeds in net production in a large individual of 20 g dry weight.     

栓皮栎叶片解剖结构种群变异及其对环境因子的适应

【目的】厘清天然广布树种栓皮栎叶片解剖结构的种群变异及其与环境因子的关系。【方法】以中国28个天然分布栓皮栎种群叶片为研究对象,用常规石蜡切片结合光学显微技术对栓皮栎叶片解剖性状进行研究,用巢式方差分析、Pearson相关性分析等方法对栓皮栎叶片解剖结构的种群变异与环境因子关系进行分析。【结果】(1)栓皮栎叶片解剖性状在28个种群间存在显著差异,平均变异系数为7.84%～15.16%,且同一叶片解剖性状在不同种群间的变异范围不同;(2)9个解剖性状在种群内及种群间均存在极显著差异,平均表型分化系数为37.44%,解剖性状变异主要来自种群内;(3)叶片厚度、上表皮厚度、栅栏组织厚度、海绵组织厚度与纬度呈显著正相关关系,年均气温及年降水量对上表皮厚度和栅栏组织厚度有显著影响。【结论】栓皮栎具有丰富的遗传多样性,为适应低温与干旱,栓皮栎叶片整体呈增厚趋势。研究结果为了解栓皮栎环境适应策略提供理论依据。     

Relative growth rate in phylogenetically related deciduous and evergreen woody species

Relative growth rate (RGR) and other growth parameters were studied in eight pairs of closely related deciduous and evergreen species (within the same genus or family). The main objective of this study was to test the association between leaf turnover rate and RGR, specific leaf area (SLA, leaf area/leaf dry weight) and other growth variables. Plants were grown for 6 months in a greenhouse under favourable water and nutrient conditions. Variation in RGR among the 16 woody species was due mainly to differences in morphological parameters such as leaf area ratio (LAR, whole plant area/whole plant dry weight) and SLA). However, temporal variation in RGR within species was due mainly to variation in net assimilation rate. When phylogeny was not taken into account, analyses showed that deciduous species grew faster than evergreens. In contrast, when phylogeny was taken into account, the data analysis showed that a faster RGR is not consistently associated with the deciduous habit (in five pairs it was, but in the other three it was not). The faster growth of the deciduous trees (in the five positive contrasts) could be explained by their higher LAR and higher SLA relative to evergreens. The lack of differences in RGR between deciduous and evergreens (in three pairs) was due to the higher leaf mass ratio (LMR, leaf dry biomass/total dry biomass) for the evergreens, which offset the higher SLA of the deciduous species, resulting in a similar LAR in both functional groups (LAR=LMR2SLA). Deciduous species had consistently higher SLA than evergreens. We suggest that SLA, more than RGR, could be an important parameter in determining adaptive advantages of deciduous and evergreen species.