干热河谷草本植物生物量分配及其对环境因子的响应

以干热河谷6种草本植物为对象,研究了水分、养分、刈割对生物量在根、茎、叶的分配及异速生长关系的影响.结果表明:刈割处理叶生物量质量分数从25.1%显著增加到31.2%,茎生物量质量分数从43.7%显著降低到34.2%;养分添加处理根生物量质量分数从34.0%显著降低到30.8%;水分处理对生物量分配没有显著影响.物种对根、茎、叶生物量分配有显著影响,适应贫瘠土壤的物种将更多生物量分配给叶和根,对茎生物量的分配相对较低.物种与环境因子存在显著的互作效应,表明环境因子对不同物种的生物量分配影响不同.适应贫瘠土壤的物种叶-茎标度指数和异速生长常数大于其他物种,而茎-根标度指数和异速生长常数小于其他物种.养分显著增加了叶-茎和叶-根的异速生长常数,刈割显著降低了茎-根的标度指数,水分处理则没有显著效应.环境因素对器官间异速生长关系的影响存在种间差异.生物量分配的种间差异及其对环境因素的响应特征可能对植物适应环境变化产生重要影响.     

Phylogenetic and growth form variation in the scaling of nitrogen and phosphorus in the seed plants

Plant biomass and nutrient allocation explicitly links the evolved strategies of plant species to the material and energy cycles of ecosystems. Allocation of nitrogen (N) and phosphorus (P) is of particular interest because N and P play pivotal roles in many aspects of plant biology, and their availability frequently limits plant growth. Here we present a comparative scaling analysis of a global data compilation detailing the N and P contents of leaves, stems, roots, and reproductive structures of 1,287 species in 152 seed plant families. We find that P and N contents (as well as N : P) are generally highly correlated both within and across organs and that differences exist between woody and herbaceous taxa. Between plant organs, the quantitative form of the scaling relationship changes systematically, depending on whether the organs considered are primarily structural (i.e., stems, roots) or metabolically active (i.e., leaves, reproductive structures). While we find significant phylogenetic signals in the data, similar scaling relationships occur in independently evolving plant lineages, which implies that both the contingencies of evolutionary history and some degree of environmental convergence have led to a common set of rules that constrain the partitioning of nutrients among plant organs.     

甘肃南部7种高寒杜鹃生物量分配的异速生长关系

生物量分配模式影响着植物个体生长和繁殖到整个群落的质量和能量流动等所有层次的功能, 揭示高寒灌丛的生物量分配模式不仅可以掌握植物的生活史策略, 而且对理解灌丛碳汇不确定性具有重要意义。该研究以甘肃南部高山-亚高山区的常绿灌丛——杜鹃(Rhododendron spp.)灌丛的7个典型种为对象, 采用全株收获法研究了不同物种个体水平上各器官生物量的分配比例和异速生长关系。结果表明: 7种高寒杜鹃根、茎、叶生物量的分配平均比例为35.57%、45.61%和18.83%, 各器官生物量分配比例的物种差异显著; 7种高寒杜鹃的叶与茎、叶与根、茎与根以及地上生物量与地下生物量之间既有异速生长关系, 也有等速生长关系, 异速生长指数不完全支持生态代谢理论和小个体等速生长理论的参考值; 各器官异速生长关系的物种差异显著。结合最优分配理论和异速生长理论能更好地解释陇南山地7种高寒杜鹃生物量的变异及适应机制。     

Linking leaf and root trait syndromes among 39 grassland and savannah species

Here, we tested hypothesized relationships among leaf and fine root traits of grass, forb, legume, and woody plant species of a savannah community. CO2 exchange rates, structural traits, chemistry, and longevity were measured in tissues of 39 species grown in long-term monocultures. Across species, respiration rates of leaves and fine roots exhibited a common regression relationship with tissue nitrogen (N) concentration, although legumes had lower rates at comparable N concentrations. Respiration rates and N concentration declined with increasing longevity of leaves and roots. Species rankings of leaf and fine-root N and longevity were correlated, but not specific leaf area and specific root length. The C3 and C4 grasses had lower N concentrations than forbs and legumes, but higher photosynthesis rates across a similar range of leaf N. Despite contrasting photosynthetic pathways and N2-fixing ability among these species, concordance in above- and below-ground traits was evident in comparable rankings in leaf and root longevity, N and respiration rates, which is evidence of a common leaf and root trait syndrome linking traits to effects on plant and ecosystem processes.     

Evolutionary size changes in plants of the south‐west Pacific

Aim To investigate evolutionary changes in the size of leaves, stems and seeds of plants inhabiting isolated islands surrounding New Zealand. Location Antipodes, Auckland, Campbell, Chatham, Kermadec, Three Kings and Poor Knights Islands. Methods First, we compared the size of leaves and stems produced by 14 pairs of plant taxa between offshore islands and the New Zealand mainland, which were grown in a common garden to control for environmental effects. Similar comparisons of seed sizes were made between eight additional pairs of taxa. Second, we used herbarium specimens from 13 species pairs to investigate scaling relationships between leaf and stem sizes in an attempt to pinpoint which trait might be under selection. Third, we used herbarium specimens from 20 species to test whether changes in leaf size vary among islands located at different latitudes. Lastly, we compiled published records of plant heights to test whether insular species in the genus Hebe differed in size from their respective subgenera on the mainland. Results Although some evidence of dwarfism was observed, most insular taxa were larger than their mainland relatives. Leaf sizes scaled positively with stem diameters, with island taxa consistently producing larger leaves for any given stem size than mainland species. Leaf sizes also increased similarly among islands located at different latitudes. Size changes in insular Hebe species were unrelated to the average size of the respective subgenera on the mainland. Main conclusions Consistent evidence of gigantism was observed, suggesting that plants do not obey the island rule. Because our analyses were restricted to woody plants, results are also inconsistent with the ‘weeds‐to‐trees’ hypothesis. Disproportionate increases in leaf size relative to other plant traits suggest that selection may favour the evolution of larger leaves on islands, perhaps due to release from predation or increased intra‐specific competition.     

Initial burst of root development with decreasing respiratory carbon cost in Fagus crenata Blume seedlings

As terrestrial plants are rooted in one place, their metabolism must be acclimatized to continuously changing environmental conditions. This process is influenced by different metabolic traits of plant organs during ontogeny. However, direct measurement of organ-specific metabolic rates is particularly scarce, and little is known about their roles in whole-plant metabolism. In this study, we investigated size scaling of respiration rate, fresh mass and surface area of leaves, stems and roots in 65 seedlings of Fagus crenata Blume (2 weeks to 16 months old). With the increase in plant mass, the proportion of roots in whole plants increased from 20.8 to 87.3% in fresh mass and from 12.8 to 95.0% in surface area, whereas there was only a 15.6 to 60.2% increase in respiration rate. As a result, the fresh-mass-specific and surface-area-specific respiration rates in the roots decreased by 85% and 90%, respectively, and these decreases were significantly size dependent. However, such a size-dependent decrease was not observed for the surface-area-specific respiration rate in the leaves and stems. It is likely that this rapid root development is specific to the early growth stage after germination and would help plants acquire water and nutrients efficiently (i.e., at relatively low respiratory carbon costs). Overall, it is probable that the establishment of F. crenata forests and survival of F. crenata seedlings could be promoted by substantial root growth, with a reduction in respiratory carbon cost.     

The relationships among root and leaf traits of 76 grassland species and relative abundance along fertility and disturbance gradients

For 76 annual, biennial, and perennial species common in the grasslands of central Minnesota, USA, we determined the patterns of correlations among seven organ-level traits (specific leaf area, leaf thickness, leaf tissue density, leaf angle, specific root length, average fine root diameter, and fine root tissue density) and their relationships with two traits relating to growth form (whether species existed for part of the growing season in basal, non-caulescent form and whether species were rhizomatous or not). The first correlation of traits showed that grasses had thin, dense leaves and thin roots while forbs had thick, low-density leaves and thick roots without any significant differences in growth form or life history. The second correlation of traits showed a gradient of species from those with high-density roots and high-density erect leaves to species with low-density roots and low-density leaves that were held parallel to the ground. High tissue density species were more likely to exist as a basal rosette for part of the season, were less likely to be rhizomatous, and less likely to be annuals. We examined the relationships between the two axes that represent the correlations of traits and previously collected data on the relative abundance of species across gradients of nitrogen addition and disturbance. Grasses were generally more abundant than forbs and the relative abundance of grasses and forbs did not change with increasing nitrogen addition or soil disturbance. High tissue density species became less common as fertility and disturbance increased.     

Shifts in diversity and community structure of endophytic bacteria and archaea across root, stem and leaf tissues in the common reed, Phragmites australis, along a salinity gradient in a marine tidal wetland of northern China

The effects of salt stress on endophytic prokaryotic communities in plants are largely unknown, and the distribution patterns of bacterial and archaeal endophytes in different tissues of a plant species are rarely compared. We investigated the endophytic bacterial and archaeal communities in roots, stems and leaves of the common reed, Phragmites australis, collected from three tidal zones along a salinity gradient, using terminal restriction fragment (T-RF) length polymorphism analysis of the 16S rRNA genes. The results showed that the bacterial diversity in the roots was significantly higher than that in the leaves, whereas similar archaeal diversity was revealed for either plant tissues or tidal zones. Network analysis revealed that T-RFs were grouped largely by tissue, and the major groups were generally linked by a few common T-RFs. Unique T-RFs in roots were mainly present in plants growing in the supratidal zone, but unique T-RFs in stems and leaves were mainly present in those from the middle and high tidal zones. Non-metric multidimensional scaling ordination and analysis of similarity revealed that bacterial communities were significantly different among tissues (P < 0.05), but similar among tidal zones (P = 0.49). However, the archaeal communities differed among tidal zones (P < 0.05), but were similar among tissues (P = 0.89). This study indicates that: (1) the endophytic archaeal communities are influenced more significantly than the endophytic bacterial communities by soil salinity, and (2) the differential distribution patterns of bacterial and archaeal endophytes in plant tissues along a salinity gradient imply that these two groups play different roles in coastal hydrophytes.     

Antibiotic‐induced effects on scaling relationships and on plant element contents in herbs and grasses

Plant performance is correlated with element concentrations in plant tissue, which may be impacted by adverse chemical soil conditions. Antibiotics of veterinary origin can adversely affect plant performance. They are released to agricultural fields via grazing animals or manure, taken up by plants and may be stored, transformed or sequestered by plant metabolic processes. We studied the potential effects of three antibiotics (penicillin, sulfadiazine, and tetracycline) on plant element contents (macro‐ and microelements). Plant species included two herb species (Brassica napus and Capsella bursa‐pastoris) and two grass species (Triticum aestivum and Apera spica‐venti), representing two crop species and two noncrop species commonly found in field margins, respectively. Antibiotic concentrations were chosen as to reflect in vivo situations, that is, relatively low concentrations similar to those detected in soils. In a greenhouse experiment, plants were raised in soil spiked with antibiotics. After harvest, macro‐ and microelements in plant leaves, stems, and roots were determined (mg/g). Results indicate that antibiotics can affect element contents in plants. Penicillin exerted the greatest effect both on element contents and on scaling relationships of elements between plant organs. Roots responded strongest to antibiotics compared to stems and leaves. We conclude that antibiotics in the soil, even in low concentrations, lead to low‐element homeostasis, altering the scaling relationships between roots and other plant organs, which may affect metabolic processes and ultimately the performance of a plant.     

Occurrence of endodermis with a casparian strip in stem and leaf

It is well known that an endodermis with casparian strip always occurs in roots, but few people are aware that it also occurs in stems and leaves of some vascular plants. The rather sparse literature on endodermis in aerial organs was last included in a review in 1943. The present compilation, which does not consider hydathodes, nectaries, or other secretory structures, emphasizes distribution of cauline and foliar endodermis with casparian strip. It occurs unevenly among major taxa: quite common in rhizomes and leaves among pteridophyte groups, with exceptions; absent in gymnosperm stems but found in leaves at least among some conifers; in stems of at least 30 mostly herbaceous angiosperm families, but far less common in leaves, where it is mostly reported from petioles. Etiolation can induce casparian strips in stems and petioles of some herbaceous plants, but results from leaf blades are questionable. There are recent reports of an endodermis with casparian strip in leaves of both woody and herbaceous taxa. The physiological function, if any, of a casparian strip in aerial organs remains unknown.     

Phenotypic correlates of the lianescent growth form: a review

Background

As proposed by Darwin, climbers have been assumed to allocate a smaller fraction of biomass to support organs in comparison with self-supporting plants. They have also been hypothesized to possess a set of traits associated with fast growth, resource uptake and high productivity.

Scope

In this review, these hypotheses are evaluated by assembling and synthesizing published and unpublished data sets from across the globe concerning resource allocation, growth rates and traits of leaves, stems and roots of climbers and self-supporting species.

Conclusions

The majority of studies offer little support for the smaller allocation of biomass to stems or greater relative growth rates in climbers; however, these results are based on small sized (<1 kg) plants. Simulations based on allometric biomass equations demonstrate, however, that larger lianas allocate a greater fraction of above-ground biomass to leaves (and therefore less biomass to stems) compared with similar sized trees. A survey of leaf traits of lianas revealed their lower average leaf mass per area (LMA), higher N and P concentration and a slightly higher mass-based photosynthetic rate, as well as a lower concentration of phenolic-based compounds than in woody self-supporting species, consistent with the specialization of lianas towards the fast metabolism/rapid turnover end of the global trait spectra. Liana stems have an efficient hydraulic design and unique mechanical features, while roots appear to penetrate deeper soil levels than in trees and are often able to generate hydraulic pressure. Much remains to be learned, however, about these and other functional specializations of their axial organs and the associated trade-offs. Developmental switches between self-supporting, searcher and climbing shoots within the same individual are a promising field of comparative studies on trait association in lianas. Finally, some of the vast trait variability within lianas may be reduced when species with different climbing mechanisms are considered separately, and when phylogenetic conservatism is accounted for.     

青藏高原三种优势植物生物量分配的变化规律

该研究利用4个由高到低不同海拔的同质园实验,以青藏高原高寒草地优势植物垂穗披碱草(Elymus nutans)、矮嵩草(Kobresia humilis)和珠芽蓼(Polygonum viviparum)为对象,分析了植物个体根、茎、叶生物量分配及根冠比的变化规律及影响因素。结果表明:(1)植物个体根、茎、叶质量比和根冠比具有显著的种间差异;与垂穗披碱草和珠芽蓼相比,矮嵩草具有显著较高的根质量比而叶、茎质量比较低,所以其根冠比较高。(2)在向低海拔移栽的过程中,珠芽蓼叶质量比保持不变,茎质量比显著降低而根质量比显著升高,根冠比表现出显著上升的趋势;垂穗披碱草则相反,即叶、茎质量比显著升高而根质量比显著降低,根冠比表现出显著下降的趋势;矮嵩草根、茎、叶质量比和根冠比则无显著变化。(3)随着海拔降低,年均气温明显升高而年均降雨量明显降低,且在植物个体种源地和土壤基质保持一致的条件下,向低海拔移栽过程中温度是导致珠芽蓼根、茎、叶生物量分配及根冠比变化的重要因素,而水分是垂穗披碱草根、茎、叶生物量分配及根冠比变化的重要驱动因素;矮嵩草根、茎、叶生物量分配及根冠比受其遗传因素影响较大。因此,在将来暖干化的背景下,青藏高原高寒草地植物生物量的分配将会发生改变,导致它们对资源(光照、水分和土壤养分)获取和利用的变化而改变它们的种间关系,从而影响群落的物种多样性与组成,最终可能导致生态系统功能的变化。     

The influence of phylogeny and ecology on root,shoot and plant ionomes of 14 native Brazilian species

The ionome is the elemental composition of a living organism, its tissues, cells or cell compartments. The ionomes of roots, stems and leaves of 14 native Brazilian forest species were characterised to examine the relationships between plant and organ ionomes and the phylogenetic and ecological affiliations of species. The null hypothesis that ionomes of Brazilian forest species and their organs do not differ was tested. Concentrations of mineral nutrients in roots, stems and leaves were determined for 14 Brazilian forest species, representing seven angiosperm orders, grown hydroponically in a complete nutrient solution. The 14 species could be differentiated by their ionomes and the partitioning of mineral nutrients between organs. The ionomic differences between the 14 species did not reflect their phylogenetic relationships or successional ecology. Differences between shoot ionomes and root ionomes were greater than differences in the ionome of an organ when compared among genotypes. In conclusion, differences in ionomes of species and their organs reflect a combination of ancient phylogenetic and recent environmental adaptations.     

Comparisons of photosynthesis-related traits among understory lichens,mosses and vascular plant leaves in a high-elevation subalpine forest

对比高海拔亚高山森林林下地衣、藓类及维管植物叶片的光合相关性状 在高海拔亚高山森林中，许多地衣和藓类植物欣欣向荣，甚至成为林地表层的优势物种。尽管它们在森林生态系统中承担着不可替代的功能，但我们对其光合作用相关的功能性状水平知之甚少，对相同生境下藓类植物和维管植物的比较研究也较为匮乏。因此，本研究选择中国青藏高原东缘亚高山林下常见的3种地衣、3种藓类和4种维管植物，测定并对比了它们的碳、氮、磷含量及化学计量比、单位面积光合组织干重、叶绿素含量及光合光响应曲线的异同。研究结果表明，林下地衣的氮含量高于藓类植物，二者的氮、磷、叶绿素、光合速率及光合氮利用效率均低于维管植物叶片。不同种类地衣的光合相关性状与其生长环境相符，而林地表层的3种藓类植物和3种草本植物的功能性状在各自功能群中的种间差异均不显著。总之，这些结果表明高海拔亚高山林下地衣和藓类植物的优势与其光合相关性状关系不大，对严酷环境较强的适应能力可能是它们得以繁荣生长的关键。     

Acclimation of Respiratory O2 Uptake in Green Tissues of Field-Grown Native Species after Long-Term Exposure to Elevated Atmospheric CO2

C3 and C4 plants were grown in open-top chambers in the field at two CO2 concentrations, normal ambient (ambient) and normal ambient + 340 [mu]LL-1 (elevated). Dark oxygen uptake was measured in leaves and stems using a liquid-phase Clark-type oxygen electrode. High CO2 treatment decreased dark oxygen uptake in stems of Scirpus olneyi (C3) and leaves of Lindera benzoin (C3) expressed on either a dry weight or area basis. Respiration of Spartina patens (C4) leaves was unaffected by CO2 treatment. Leaf dry weight per unit area was unchanged by CO2, but respiration per unit of carbon or per unit of nitrogen was decreased in the C3 species grown at high CO2. The component of respiration in stems of S. olneyi and leaves of L. benzoin primarily affected by long-term exposure to the elevated CO2 treatment was the activity of the cytochrome pathway. Elevated CO2 had no effect on activity and capacity of the alternative pathway in S. olneyi. The cytochrome c oxidase activity, assayed in a cell-free extract, was strongly decreased by growth at high CO2 in stems of S. olneyi but it was unaffected in S. patens leaves. The activity of cytochrome c oxidase and complex III extracted from mature leaves of L. benzoin was also decreased after one growing season of plant exposure to elevated CO2 concentration. These results show that in some C3 species respiration will be reduced when plants are grown in elevated atmospheric CO2. The possible physiological causes and implications of these effects are discussed.     

4种鼠尾草属植物根、茎、叶中脂溶性成分的分布差异性

为了说明4种鼠尾草属植物脂溶性成分在根、茎、叶中的分布情况,本文采用了组织化学定位和HPLC图谱分析相结合的方法,对4种鼠尾草属植物不同部位的脂溶性成分进行了分析.结果表明:4种鼠尾草属植物的根、茎、叶中脂溶性成分的分布有差异,根中脂溶性成分分布于周皮,茎、叶中脂溶性成分分布于表皮.丹参和白花丹参根中的脂溶性成分主要是丹参酮ⅡA、丹参酮Ⅰ和二氢丹参酮Ⅰ,鼠尾草和药用鼠尾草根中仅含丹参酮ⅡA,不含丹参酮Ⅰ和二氢丹参酮Ⅰ.茎、叶中的脂溶性成分不是丹参酮ⅡA、丹参酮Ⅰ和二氢丹参酮Ⅰ.脂溶性成分的种类和含量随品种和引种地的不同而产生差异.这一结果明确了脂溶性成分在4种鼠尾草属植物不同部位的确切分布,为准确用药提供了依据.     

Leaf position,light levels,and nitrogen allocation in five species of rain forest pioneer trees

We used path analysis to ask whether leaf position or leaf light level was a better predictor of within-plant variation in leaf nitrogen concentration in five species of rain forest pioneer trees (Cecropia obtusifolia, Ficus insipida, Heliocarpus appendiculatus, Piper auritum, and Urera caracasana) from the Los Tuxtlas Biological Station, Veracruz, Mexico. Three hundred seventy-five leaves on 28 plants of the five species were analyzed for leaf nitrogen concentration, leaf mass per area, and leaf light interception at different positions (= nodes) along a shoot. Mean values of leaf nitrogen concentration ranged from 0.697 to 0.993 g/m² in the five species, and varied by as much as 2.24 g/m² among leaves on individual plants. Leaf position on the shoot explained significantly more of the within-plant variation in leaf nitrogen concentration than did leaf light level in four of the five species: Cecropia obtusifolia, Heliocarpus appendiculatus, Piper auritum (branch leaves only), and Urera caracasana. However, individual species differed considerably in the patterns of nitrogen allocation and leaf mass per area among leaves on a shoot. These results suggest that leaf nitrogen deployment in these plants is, in part, developmentally constrained and related to the predictability of canopy light distribution associated with plant growth form.     

超富集植物短毛蓼对锰的富集特征

通过野外调查和营养液培养试验,研究了锰在短毛蓼体内的富集特征和对其生长的影响.在锰含量高达2.5×105mg/kg的锰矿废弃地上短毛蓼生长良好,叶锰含量高达1.66×104mg/kg.营养液培养条件下,随着生长介质中Mn浓度的升高,短毛蓼根、茎、叶中的Mn含量逐渐增加,当锰供应水平为1.000mmol/L时,叶锰含量超过10000mg/kg;当锰供应水平为20 000mmol/L时,短毛蓼仍能生长,根、茎和叶3部分的锰含量均达到最大值,分别为9923,18112mg/kg和55750mg/kg.在所有锰供应水平下,短毛蓼茎和叶中的锰含量都比根部的高.结果表明,短毛蓼是一种锰超富集植物,这一发现为锰污染土壤的植物修复和探讨锰在植物体内的超富集机理提供了一种新的种质资源.     

Stem hypoxia and root respiration of flooded maple and birch seedlings

Some researchers have attributed flood tolerance of woody plants to air entering the shoot through stems, leaves, or lenticels and diffusing to the roots to sustain aerobie respiration. The purpose of this study was to determine if internal aeration of roots by lower stems, changes in gross morphology of lower stems, or both, contribute to flood tolerance of certain tree species. Greenhouse-grown seedlings of red maple ( Acer rubrum L.) and river birch ( Betula nigra L.) tolerated at least 30 days of flooding, where as sugar maple ( Acer saccharum Marsh) and European white birch (also called silver birch, Betula pendula Roth) were intolerant. Flood treatment induced lentieel intumescences and adventitious root formation on red maple stems, but only adventitious roots formed on river birch stems. Stem morphology of sugar maple and European birch was unchanged by flooding. Flood stress decreased oxygen consumption capacity of excised roots from both tolerant and intolerant species. Exclusion of oxygen from the lower stems of flooded red maple and river birch prevented lenticel intumescence and adventitious root formation, but flood tolerance and root respiration capacity were unchanged. Neither internal aeration nor changes in stem morphology appear to account for flood tolerance of red maple and river birch.     

Stem and wood allometric relationships in Cacteae (Cactaceae)

Allometric relationships in organisms are considered a universal phenomenon. A positive scaling has been reported between stem size and cellular size of tracheary elements in wood of different vascular plants, but few studies have been carried out in slow-growing succulent plants. The aim of this study was to evaluate if a relationship exists between size, growth form and wood cell size among individual species of Cacteae. Forty-four species belonging to 16 genera of the tribe Cacteae with differing growth forms and sizes were studied. When analyzing plant size, we found a positive allometric scaling and the larger-sized species showing a higher percentage of succulent tissue and less accumulation of wood tissue. The positive scaling found between plant size (height and diameter) and vessel elements and fiber length support the universality of the allometric relationship proposed for other vascular plants with non-succulent stems. Notably, wide-band tracheids do not scale with plant size or growth form. Succulence associated with narrow vessel elements with distinctive helical secondary walls and wide-band tracheids suggest they are the key adaptations to tolerate drought and provide support to the stems of most taxa in Cacteae. Fibers do not have the primary role of giving mechanical support; therefore, we assume the scarce fibers in clusters represent reaction wood that, along with the fundamental tissue, maintains the vertical position and shape of those species growing in rocky cracks. Our results with species having short succulent stems support the universal theory of positive allometric scaling of vascular plants.