The leaf size-twig size spectrum of temperate woody species along an altitudinal gradient: an invariant allometric scaling relationship

BACKGROUND AND AIMS: The leaf size-twig size spectrum is one of the leading dimensions of plant ecological variation, and now it is under development. The purpose of this study was to test whether the relationship between leaf size and twig size is isometric or allometric, and to examine the relationship between plant allometric growth and life history strategies in the spectrum. METHODS: Leaf and stem characters-including leaf and stem mass, total leaf area, individual leaf area, stem cross-sectional area, leaf number and stem length-at the twig level for 59 woody species were investigated along an altitudinal gradient on Changbaishan Mountain in the temperate zone of China. The environmental gradient ranges from temperate broad-leaved mixed forest at low altitude, to conifer forest at middle altitude, and to sub-alpine birch forest at high altitude. The scaling relationships between stem cross-sectional area and stem mass, stem mass and leaf mass, and leaf mass and leaf area at the twig level were simultaneously determined. KEY RESULTS: Twig cross-sectional area was found to have invariant allometric scaling relationships with the stem mass, leaf mass, total leaf area and individual leaf area, all with common slopes being significantly larger than 1, for three altitudinal-zoned vegetation types under investigation. However, leaf mass was found to be isometrically related to stem mass and leaf area along the environmental gradient. Based on the predictions of previous models, the exponent value of the relationship between twig cross-sectional area and total leaf area can be inferred to be 1.5, which falls between the confidence intervals of the relationship at each altitude, and between the confidence intervals of the common slope value (1.17-1.56) of this study. This invariant scaling relationship is assumed to result from the fractural network and/or developmental constraints of plants. The allometric constants (y-intercepts) of the relationships between the stem cross-sectional area and leaf area (both total leaf area and individual leaf area) were found to decrease significantly along the altitudinal gradient. This suggests that the species would support less leaf area at a given twig cross-sectional area with increasing environmental stress. CONCLUSIONS: This study demonstrated that plants respond to the environmental gradient by changing the y-intercepts of the relationship between leaf size-twig size, while keeping the exponent value of the allometric relationship as an invariant constant. The allometric growth in the twig size-leaf size spectrum is related to many other components of plant life history strategy, including the well established life history trade-off between efficiency and safety in the hydraulic transport of water.     

Reconciling the apparent difference between mass- and area-based expressions of the photosynthesis-nitrogen relationship

The relationship between photosynthetic carbon assimilation (A _max) and leaf nitrogen content (N _leaf) can be expressed on either a leaf area basis (A _area vs N _area) or a leaf mass basis (A _mass vs N _mass). Dimensional analysis shows that the units for the slope of this relationship are the same for both expressions (μmol [CO₂] g⁻¹ [N] s⁻¹). Thus the slope measures the change in CO₂ assimilation per gram of nitrogen, independent of leaf mass or leaf area. Although they have the same units, large differences between the area and mass-based slopes have been observed over a broad range of taxonomically diverse species. Some authors have claimed that regardless of these differences, the fundamental nature of the A _max-N _leaf relationship is independent of the units of expression. In contrast, other authors have claimed that the area-based A _max-N _leaf relationship is fundamentally different from the mass-based relationship because of interactions between A _max, N _leaf, and leaf mass per area (LMA, g [leaf] m⁻² [leaf]). In this study we consider the mathematical relationships involved in the transformation from mass- to area-based expressions (and vice versa), and the implications this transformation has for the slope of the A _max-N _leaf relationship. We then show that the slope of the relationship is independent of the units of expression when the effect of LMA is controlled statistically using a multiple regression. The validity of this hypothesis is demonstrated using 13 taxonomically and functionally diverse C₃ species. This analysis shows that the slope of the A _max-N _leaf relationship is similar for the mass- and area-based expressions and that significant errors in the estimate of the slope can arise when the effect of LMA is not controlled. Received: 7 May 1998 / Accepted: 19 October 1998     

三个东方百合品种在滇中鳞茎繁育中的光合特性

对西伯利亚、索尔邦、蒂伯3个东方百合品种(Lilium“Oriental Hybrids”)鳞茎繁育过程中植株的气体交换、叶氮含量、叶绿素含量等进行了测定,试图了解东方百合在滇中低纬高原地区的生理生态适应性。3个品种中,蒂伯的光合能力最强,西伯利亚最弱,其光合速率与羧化效率、比叶重及叶氮含量相关,说明其光合能力主要受Rubisco的活性和/或数量影响。3个品种的光补偿点和光饱和点较低,且在高光下没有明显的光抑制现象,对光强表现出宽广的适应性。不同品种的光合最适温度不同,西伯利亚较高(25.5～34.9℃),适宜种植于较温暖的地区;索尔邦较低(19.3～25.6℃),适宜种植于冷凉地区;蒂伯对温度较不敏感,在滇中低纬高原的气候条件下有更广的种植区域。     

Anatomical basis of the change in leaf mass per area and nitrogen investment with relative irradiance within the canopy of eight temperate tree species

Changes in leaf mass per area (LMA), nitrogen content on a mass-basis (N_m) and on an area basis (N_a) with relative irradiance were assessed in leaves of eight temperate species harvested at different depths in a canopy. Relative irradiance (GSF) at the points of leaf sampling was estimated by hemispheric photographs. There was a strong species-dependent positive relationship between LMA and GSF for all species. Shade-tolerant species such as Fagus sylvatica showed lower LMA for the same GSF than less tolerant species as Quercus pyrenaica or Quercus petraea. The only evergreen species in the study, Ilex aquifollium, had the highest LMA, independent of light environment, with minimum values much higher than the rest of the broad-leaved species studied. There was no relation between N_m and GSF for most species studied and only a very weak relation for the relative shade-intolerant species Q. pyrenaica. Within each species, the pattern of N_a investment with regard to GSF was linked mainly to LMA. At the same relative irradiance, differences in N_a among species were conditioned both by the LMA–GSF relationship and by the species N_m value. The lowest N_m value was measured in I. aquifollium (14.3 ± 0.6 mg g^–1); intermediate values in Crataegus monogyna (16.9 ± 0.6 mg g^–1) and Prunus avium (19.1 ± 0.6 mg g^–1) and higher values, all in a narrow range (21.3 ± 0.6 to 23 ± 0.6 mg g^–1), were measured for the other five species. Changes in LMA with the relative irradiance were linked both to lamina thickness (LT) and to palisade/spongy parenchyma ratio (PP/SP). In the second case, the LMA changes may be related to an increase in lamina density as palisade parenchyma involves higher cell packing than spongy parenchyma. However, since PP/SP ratio showed a weak species-specific relationship with LMA, the increase in LT should be the main cause of LMA variation.     

Plant growth-form alters the relationship between foliar morphology and species shade-tolerance ranking in temperate woody taxa

Variation in leaf size (area per leaf) and leaf dry weight per area (LWA) in relation to species shade- and drought-tolerance, characterised by Ellenberg's light (ELD) and water demand (EWD) values, respectively, were examined in 60 temperate woody taxa at constant relative irradiance. LWA was independent of plant size, but leaf size increased with total plant height at constant ELD. Canopy position also affected leaf morphology: leaves from the upper crown third had higher LWA and were larger than leaves from the lower third. Leaf size and LWA were negatively correlated, and leaf size decreased and LWA increased with decreasing species shade-tolerance. Mean LWA was similar for trees and shrubs, but trees had larger leaves than shrubs. Furthermore, all relationships were altered by plant growth-form: none of the qualitative tendencies was significant for trees. This implies the considerably lower plasticity of foliar parameters in trees than those in shrubs. Accordingly, shade-tolerance of trees, having relatively constant leaf structure, may be most affected by the variability in biomass partitioning and crown geometry which influence foliage distribution and spacing and finally determine canopy light absorptance. Alteration of leaf form and investment pattern for construction of unit foliar surface area which change the efficiency of light interception per unit biomass investment in leaves, is a competitive strategy inherent to shrubs. EWD as well as wood anatomy did not control LWA and leaf size, though there was a trend of ring-porous tree species to be more shade-tolerant than diffuse-porous trees. Since ring-porous species are more vulnerable to cavitation than diffuse-porous species, they may be constrained to environments where irradiances and consequently evaporative demand is lower.     

Scaling of offspring number and mass to plant and animal size: model and meta-analysis

The scaling of reproductive parameters to body size is important for understanding ecological and evolutionary patterns. Here, we derived allometric relationships for the number and mass of seeds, eggs and neonates from an existing model on population production. In a separate meta-analysis, we collected 79 empirical regressions on offspring mass and number covering different taxa and various habitats. The literature review served as a validation of the model, whereas, vice versa, consistency of isolated regressions with each other and related ecological quantities was checked with the model. The total offspring mass delivered in a reproductive event scaled to adult size with slopes in the range of about 3/4 to 1. Exponents for individual seed, egg and neonate mass varied around 1/2 for most heterotherms and between 3/4 and 1 for most homeotherms. The scaling of the progeny number released in a sowing, clutch or litter was opposite to that of their size. The linear regressions fitted into a triangular envelope where maximum offspring mass is limited by the size of the adult. Minimum seed and egg size scaled with weight exponents of approximately 0 up to 1/4. These patterns can be explained by the influence of parents on the fate of their offspring, covering the continuum of r-strategists (pelagic–aquatic, arial, most invertebrates, heterotherms) and K-strategists (littoral–terrestrial, some invertebrates, homeotherms).     

Root hydraulic conductivity and whole-plant water balance in tropical saplings following a shade-to-sun transfer

We hypothesized that pioneer and late successional species show different morphological and physiological responses in water use after gap formation. The magnitude of the responses was compared between two pioneer species (Macaranga gigantea and Trema orientalis) and four late successional species (Shorea sp.), in an experiment in which saplings were transferred from shade to sun. Although transpiration demand increased following the transfer, root hydraulic conductivity (Lp_r) decreased. Lp_r was sensitive to brief treatments with HgCl₂ (a specific inhibitor of aquaporins). This allows Lp_r to be divided into two components: cell-to-cell and apoplastic pathways. The Lp_r of cell-to-cell pathway decreased in all species following the transfer, relating to aquaporin depression in roots. Following the transfer, leaf osmotic potentials at full hydration decreased and both leaf mass per area [leaf mass/leaf area (LMA)] and fine-root surface area/leaf surface area (root SA/leaf SA) increased in almost all species, allowing saplings to compensate for the decrease in Lp_r. Physiologically, pioneer species showed larger decreases in Lp_r and more effective osmotic adjustment than late successional species, and morphologically, pioneer species showed larger increases in root SA/leaf SA and LMA. Water balance at the whole-plant level should be regulated by coupled responses between the aboveground and the belowground parts. Interspecific differences in responses after gap formation suggest niche differentiation in water use between pioneer and late successional species in accordance with canopy-gap size.     

Leaf nitrogen distribution in relation to leaf age and photon flux density in dominant and subordinate plants in dense stands of a dicotyledonous herb

We studied the effects of photon flux density (PFD) and leaf position, a measure of developmental age, on the distribution of nitrogen content per unit leaf area (N _area) in plants of different heights, in dense stands grown at two nitrogen availabilities and in solitary plants of the erect dicotyledonous herb Xanthium canadense. Taller more dominant plants received higher PFD levels and experienced a larger difference in relative PFD between their youngest and oldest leaves than shorter subordinate plants in the stands. Differences in PFD between leaves of solitary plants were assumed to be minimal and differences in leaf traits, found for these plants, could thus be mainly attributed to an effect of leaf position. In the solitary plants, N _area decreased with leaf position while in the plants from the stands it decreased with decreasing relative PFD, indicating both factors to be important in determining the distribution of N _area. Due to the effect of leaf position on N _area, leaves of subordinate plants had a higher N _area than older leaves of dominant plants which were at the same height or slightly higher in the canopy. Consequently, the N _area distribution patterns of individual plants plotted as a function of relative PFD were steeper, and probably closer to the optimal distribution which maximizes photosynthesis, than the average distribution in the stand. Leaves of subordinate plants had a lower mass per unit area (LMA) than those of dominant plants. In the dominant plants, LMA decreased with decreasing relative PFD (and with leaf position) while in the subordinate plants it increased. This surprising result for the subordinate plants can be explained by the fact that, during the course of a growing season, these plants became increasingly shaded and newer leaves were thus formed at progressively lower light availability. This indicates that LMA was strongly determined by the relative PFD at leaf formation and to a lesser extent by the current PFD. Leaf N content per unit mass (N _mass) was strongly determined by leaf position independent of relative PFD. This indicates that N _mass is strongly ontogenetically related to the leaf-aging process while changes in N _area, in response to PFD, were regulated through changes in LMA. Received: 11 May 1997 / Accepted: 9 September 1997     

Biological spectrum and other structural functional attributes of the vegetation of Kumaun Himalaya

Spectra on life form, leaf size, leaf persistence, flowering season, and shade tolerance of trees in different vegetation types occurring within the north-western catchment of the river Gola in Kumaun Himalaya are presented. The flora of Quercus leucotrichophora, Quercus lanuginosa and Quercus floribunda forests is phanerophytic, that of Pinus roxburghii and mixed forests therophytic. The grassland vegetation is characterized by the largest percentage of hemicryptophytes. The flora of the whole area, is therophytic. The biological spectrum for the entire Kumaun Himalaya is characterized as therohemigeophytic. Among the various altitudinal zones, the tropical has a preponderance of phanerophytes, the temperate and the alpine of hemicryptophytes.The observations on leaf size indicate that with the exception of Pinus roxburghii forest, in all vegetation types, the species with microphylls are greater in number. In this region, the vegetation expression is evergreen, although the tree flora has a considerable content of deciduous elements. In all the forests, the flowering period in most of the trees is vernal.On the basis of relative density, the greatest proportions of adult trees in the Pinus roxburghii and Quercus lanuginosa forests are shade intolerant, while in the mixed and Quercus leucotrichophora forests maximum trees are intermediate in shade tolerance. With the exception of the Pinus roxburghii forest, all the forests exhibit the dominance of trees which are shade tolerant at the seedling stage. On the basis of relative density, all forest types, except for Pinus roxburghii forest, have 74.5 to 100% trees with the potentiality of vegetative reproduction.Financial support from the Indian Space Research Organisation, Banglore and University Grants Commission, New Delhi is gratefully acknowledged.     

Genetic variation in leaf pigment,optical and photosynthetic function among diverse phenotypes of <Emphasis Type="Italic">Metrosideros polymorpha</Emphasis> grown in a common garden

Coordinated variation has been reported for leaf structure, composition and function, across and within species, and theoretically should occur across populations of a species that span an extensive environmental range. We focused on Hawaiian keystone tree species Metrosideros polymorpha, specifically, 13-year old trees grown (2–4 m tall) in a common garden (approximately 1 ha field with 2–3 m between trees) from seeds collected from 14 populations along an altitude–soil age gradient. We determined the genetic component of relationships among specific leaf area (SLA), the concentrations of nitrogen (N) and pigments (chlorophylls, carotenoids, and anthocyanins), and photosynthetic light-use efficiency. These traits showed strong ecotypic variation; SLA declined 35% with increasing source elevation, and area-based concentrations of N, Chl a + b and Car increased by 50, 109 and 96%, respectively. Concentrations expressed on a mass basis were not well related to source elevation. Pigment ratios expressed covariation that suggested an increased capacity for light harvesting at higher source elevation; Chl/N and Car/Chl increased with source elevation, whereas Chl a/b declined; Chl a/b was higher for populations on younger soil, suggesting optimization for low N supply. Parallel trends were found for the photosynthetic reactions; light-saturated quantum yield of photosystem II (Φ _PSII) and electron transport rate (ETR) increased with source elevation. Correlations of the concentrations of photosynthetic pigments, pigment ratios, and photosynthetic function across the ecotypes indicated a stoichiometric coordination of the components of the light-harvesting antennae and reaction centers. The constellation of coordinated morphological, biochemical and physiological properties was expressed in the leaf reflectance and transmittance properties in the visible and near-infrared wavelength region (400–950 nm), providing an integrated metric of leaf status among and between plant phenotypes.     

肉豆蔻科国家重点保护植物叶片功能性状的研究

为探究重点保护植物在热带地区的生态适应策略,本研究以肉豆蔻科9种木本植物（5种国家重点保护植物和4种非国家重点保护植物）为研究对象,测定了它们的叶片形态、光合、养分含量及养分重吸收率等22个性状。结果显示：与非重点保护植物相比,重点保护植物的叶片干物质含量、叶片密度和碳含量更低,但叶片更厚、单位叶面积光合速率更高;比叶重、叶脉密度、气孔导度、元素含量、养分重吸收率等多个性状在重点和非重点保护植物间差异不显著。氮含量与氮重吸收率显著正相关。镁含量与比叶重在重点保护植物间显著负相关,镁重吸收率与叶脉密度在重点保护与非重点保护植物间均显著负相关。主成分分析结果表明,对第1主成分贡献较大的性状主要是叶片密度、光合氮利用效率和叶片干物质含量;对第2主成分贡献较大的性状主要是N、P含量和N : P比值。肉豆蔻科植物叶片的N : P比值低于14,说明其生长主要是受N限制而不是P限制。研究结果表明,基于功能性状的研究可以揭示重点保护植物和非重点保护植物间生态适应策略的差异。因此,未来需加强重点和非重点保护植物的生理生态学比较研究。     

Variations and interrelationships of foliar hydraulic and photosynthetic traits for Larix gmelinii北大核心CSCD

Aims: Variations and potential trade-offs of leaf hydraulic and photosynthetic traits are essential for assessing and predicting the effect of climate change on tree survival, growth and distribution. Our aims were to examine variations and interrelationships of leaf hydraulic and photosynthetic traits in response to changes in site conditions for Dahurian larch (Larix gmelinii)-a dominant tree species in Chinese boreal forests. Methods: This study was conducted at the Maoershan Forest Ecosystem Research Station. A transect of 27 year-old Dahurian larch plantation was established that consisted of five plots extending from the valley to the ridge of a slope. The predawn leaf water potential (ψpre), area- and mass-based leaf hydraulic conductance (Karea and Kmass, respectively), resistance to embolism capacity (P50), leaf mass per area (LMA), net photosynthetic rate (A), and leaf nitrogen content (N) were measured in August 2016. Important findings: The ψpre, Karea, Kmass, P50, A, LMA, and N all varied significantly among the plots (p < 0.05), indicating significant intra-specific variations in these traits in response to the changes in site conditions. The P50 was significantly (p < 0.05) correlated with ψpre, Karea or Kmass, suggesting that a trade-off between hydraulic efficiency and safety exist within the species to some degree. There were significant (p < 0.05) pairwise correlations between A, LMA, and N. Nevertheless, there was no significant (p < 0.05) correlation between the measured photosynthetic traits and hydraulic traits. We concluded that the intra-specific variations and multiple interrelationships of the leaf hydraulic and photosynthetic traits for the larch reflect the plasticity of its leaf traits and strategies of its survival and growth as a result of its acclimation to diverse site conditions.     

南亚热带人工林16种木本植物重要叶特征参数的相互关系

以南亚热带退化丘陵生态恢复进程中16种3 a生木本植物为对象,研究叶片重要特征参数间的相互关系.结果表明,植物叶片的N、P含量平均值低于全国平均值,N:P平均值则高于全国的平均水平;成熟叶片的N和P含量之间有极显著正相关关系(P<0.00001),N:P与N有弱的正相关关系(r=0.322,P<0.01),与P显著负相关,推断植物叶N:P主要由P决定(r=0/639,P<0.00001).含N量高的植物具有高的最大净光合速率(Pmax)、光合N利用效率(PNOE)和低的比叶重(LMA),反之,含N量低植物具有低的Pmax、PNUE和高的LMA;Pmax与PNUE极显著正相关,LMA与PNUE极显著负相关.