Relationships of leaf dark respiration with light environment and tissue nitrogen content in juveniles of 11 cold-temperate tree species

It has been argued that plants adapted to low light should have lower carbon losses via dark respiration (Rd) than those not so adapted, and similarly, all species would be expected to down-regulate Rd in deep shade, because the associated advantages of high metabolic potential cannot be realized in such habitats. In order to test these hypotheses, and to explore the determinants of intraspecific variation in respiration rates, we measured Rd, leaf mass per unit area (LMA), and nitrogen content of mature foliage in juveniles of 11 cold-temperate tree species (angiosperms and conifers), growing in diverse light environments in forest understories in northern Minnesota. Among the seven angiosperm species, respiration on mass, area, and nitrogen bases showed significant negative overall relationships with shade tolerance level. Mass-based respiration rates (Rd _mass) of angiosperms as a group showed a significant positive overall relationship with an index of light availability (percentage canopy openness, %CO). Rd _mass of most conifers also showed evidence of acclimation of Rd _mass to light availability. LMA of all species also increased with increasing %CO, but this response was generally much stronger in angiosperms than in conifers. As a result, the response of area-based respiration (Rd _area) to %CO was dominated by ΔRd _mass for conifers, and by ΔLMA for most angiosperms, i.e., functional types differed in the components of acclimation of Rd _area to light availability. Among the seven angiosperm species, the relationships of leaf N on a mass basis (N _mass) with %CO were modulated by shade tolerance: negative slopes in shade-tolerant species may be related to the steep increases in LMA of these taxa along gradients of increasing light intensity, and associated dilution of N-rich, metabolically active tissue by increasing investment in leaf structural components. Although N _mass was therefore an unreliable predictor of variation in Rd _mass along light gradients, respiration per unit leaf N (Rd/N) was significantly positively correlated with %CO for most species. This probably reflects variation in the proportion of leaf N allocated to protein and/or the influence of leaf carbohydrate status on Rd. Species shade tolerance differences were not significantly correlated with the magnitude of either ΔRd _mass or ΔRd _area, indicating that variation in acclimation potential of Rd is much less important than inherent differences in this trait. Acclimation of Rd _mass to light availability appears to be a generalized feature of juvenile trees, and the important ecological trade-off is likely between high metabolic capacity in high light and low respiratory losses in low light. Received: 15 April 1999 / Accepted: 24 October 1999     

Modulation of leaf economic traits and trait relationships by climate

Aim Our aim was to quantify climatic influences on key leaf traits and relationships at the global scale. This knowledge provides insight into how plants have adapted to different environmental pressures, and will lead to better calibration of future vegetation–climate models. Location The data set represents vegetation from 175 sites around the world. Methods For more than 2500 vascular plant species, we compiled data on leaf mass per area (LMA), leaf life span (LL), nitrogen concentration (N_mass) and photosynthetic capacity (A_mass). Site climate was described with several standard indices. Correlation and regression analyses were used for quantifying relationships between single leaf traits and climate. Standardized major axis (SMA) analyses were used for assessing the effect of climate on bivariate relationships between leaf traits. Principal components analysis (PCA) was used to summarize multidimensional trait variation. Results At hotter, drier and higher irradiance sites, (1) mean LMA and leaf N per area were higher; (2) average LL was shorter at a given LMA, or the increase in LL was less for a given increase in LMA (LL–LMA relationships became less positive); and (3) A_mass was lower at a given N_mass, or the increase in A_mass was less for a given increase in N_mass. Considering all traits simultaneously, 18% of variation along the principal multivariate trait axis was explained by climate. Main conclusions Trait‐shifts with climate were of sufficient magnitude to have major implications for plant dry mass and nutrient economics, and represent substantial selective pressures associated with adaptation to different climatic regimes.     

漓江水陆交错带植物叶性状对水淹胁迫的响应及经济谱分析

以漓江水陆交错带为研究区,分两个条带分别量测了适生植物的5个叶性状指标:最大净光合速率(A_(max))、比叶重(LMA)、单位质量叶片全氮含量(N_(mass))、单位质量叶片全磷含量(P_(mass))、单位质量叶片全钾含量(K_(mass))。研究重度淹没带与微度淹没带不同功能型植物叶性状间的差异,分析并讨论重度淹没带叶性状间的关系与全球尺度是否存在差异,探究重度淹没带植物对水淹生境的生理响应机制。结果如下:(1)重度淹没带植物叶片的A_(mass)、N_(mass)、P_(mass)显著高于微度淹没带。(2)乔木、灌木叶片的LMA均显著高于草本植物,而A_(mass)、PPUE均显著低于草本植物。(3)重度淹没带草本叶性状指标的N_(mass)、P_(mass)、PNUE均显著高于微度微度淹没带,而乔木、灌木的叶性状在两个条带的差异则不显著。(4)重度淹没带植物叶性状关系与全球尺度基本一致,其植物叶片具有低LMA,高A_(mass)、Nmas s、P_(mass)。分析可知,重度淹没带植物在出露期提高叶片光合效率及相关营养水平可能是其适应水淹胁迫特殊生境的关键策略之一;不同功能型植物对同一环境的适应能力存在一定的差异,草本对于水淹环境的响应更为积极,适应能力更好;重度淹没带也存在叶经济谱,其植物在经济谱中属于"快速投资-收益"型物种。     

混交比例对桉树-乡土树种混交林优势树种叶片资源获取性状的影响

桉树-乡土树种混交林在提高林分生产力和生态系统功能等方面具有较大潜力。该研究以南亚热带4种桉树-乡土树种混交林(桉树与乡土树种混交比例分别为5:5、6:4、7:3、8:2)和桉树纯林为研究对象,研究了3种优势乡土树种华润楠(Machilus chinensis)、阴香(Cinnamomum burmannii)、灰木莲(Manglietia glauca)和速生树种尾叶桉(Eucalyptus urophylla)的叶片生理、结构和化学性状在不同比例混交林中的差异。结果表明,4优势造林树种的叶片性状存在明显的种间差异,其中灰木莲的比叶面积(SLA)、光合磷利用效率(PPUE)、单位质量叶片最大光合速率(A_mass)和蒸腾速率(T_mass)以及叶片养分含量最高,说明灰木莲采取资源获取型的生态策略;尾叶桉的SLA、A_mass、T_mass及叶片养分含量最低,但具有最高的PPUE,说明尾叶桉兼顾了资源获取型和保守型的物种特征。灰木莲与尾叶桉在SLA、A_mass、T_mass     

Leaf trait co‐variation,response and effect in a chronosequence

Question: Is there any generality in terms of leaf trait correlations and the multiple role of leaf traits (response to and/or effect on) during secondary succession? Location: A secondary successional sere was sampled at four different ages since abandonment from several years to nearly 150 years on the Loess Plateau of northwestern China. Method: Specific leaf area (SLA), leaf mass per area (LMA), leaf nitrogen (N_mass, N_area), leaf phosphorus (P_mass, P_area) and leaf dry matter content (LDMC) were measured for all species recorded in the successional sere. Above‐ground net primary productivity (ANPP) and specific rate of litter mass loss (SRLML) were measured as surrogates for ecosystem properties. Soil total carbon (C) and nitrogen (N) were measured in each stage. Leaf traits were related to ecosystem properties and soil nutrient gradients, respectively. Results: LMA is correlated with N_area and P_area' and negatively with N_mass. Correlation between N_area and P_area was higher than between N_mass and P_mass. At the community level, field age, community hierarchy and their interaction explain 64.4 ‐ 93.5% of the variation in leaf traits. At the species level, field age explains 22.4 ‐ 45.5% of the variation in leaf traits (excl. P_area) while plant functional group has a significant effect only for N_mass. LDMC is correlated with ANPP and negatively with SRLML; P_mass is correlated with SRLML. Conclusions: Mean values of LMA, N_mass and N_area are close to the worldwide means, suggesting that large‐scale climate has a profound effect on leaf mass and leaf nitrogen allocation, while environmental gradients represented by succession have little influence on leaf‐trait values. Correlations between leaf traits, such as LMA‐N_area, LMA‐P_area and LMA‐N_mass shown in previous studies, are confirmed here. Although none of the leaf traits is proved to be both a response trait and an effect trait independent of time scale and community hierarchy, mass‐based leaf N is likely a sensitive response trait to soil C and N gradients. In addition, LDMC can be a marker for ANPP and SRLML, while mass‐based leaf P can be a marker for SRLML.     

Biogeographic constraints on the world‐wide leaf economics spectrum

Aim The world‐wide leaf economic spectrum (LES) describes tight coordination of leaf traits across global floras, reported to date as being largely independent of phylogeny and biogeography. Here, we present and test an alternative, historical perspective that predicts that biogeography places significant constraints on global trait evolution. These hypothesized constraints could lead to important deviations in leaf trait relationships between isolated floras that were influenced by different magnitudes of genetic constraint and selection. Location Global, including floristic regions of the Northern and Southern Hemispheres, eastern North America, East Asia (EAS), the Hawaiian Islands and tropical mainland floras. Methods We use a large leaf‐trait database (GLOPNET) and species native distribution data to test for variation in leaf trait relationships modulated by floristic region, controlling for climatic differences. Standardized major axis analyses were used to evaluate biogeographic effects on bivariate relationships between LES traits, including relationships of photosynthetic capacity and dark respiration rate (A_mass–R_d‐mass), leaf lifespan and mass per area ratio (LL–LMA), and photosynthetic capacity and nitrogen content (A_mass–N_mass). Results Independent of climate or biome, floras of different evolutionary histories exhibited different leaf trait allometries. Floras of the Northern Hemisphere exhibited greater rates of return on resource investment (steeper slopes for the trait relationships analysed), and the more diverse temperate EAS flora exhibited greater slopes or intercepts in leaf trait relationships, with the exception of the A_mass–N_mass relationship. In contrast to our hypothesis, plants of the floristically isolated Hawaiian Islands exhibited a similar A_mass–N_mass relationship to those of mainland tropical regions. Main conclusions Differences in leaf trait allometries among global floristic regions support a historical perspective in understanding leaf trait relationships and suggest that independent floras can exhibit different tradeoffs in resource capture strategies.     

Relationships of leaf dark respiration to leaf nitrogen, specific leaf area and leaf life-span: a test across biomes and functional groups

Based on prior evidence of coordinated multiple leaf trait scaling, we hypothesized that variation among species in leaf dark respiration rate (R _d) should scale with variation in traits such as leaf nitrogen (N), leaf life-span, specific leaf area (SLA), and net photosynthetic capacity (A _max). However, it is not known whether such scaling, if it exists, is similar among disparate biomes and plant functional types. We tested this idea by examining the interspecific relationships between R _d measured at a standard temperature and leaf life-span, N, SLA and A _max for 69 species from four functional groups (forbs, broad-leafed trees and shrubs, and needle-leafed conifers) in six biomes traversing the Americas: alpine tundra/subalpine forest, Colorado; cold temperate forest/grassland, Wisconsin; cool temperate forest, North Carolina; desert/shrubland, New Mexico; subtropical forest, South Carolina; and tropical rain forest, Amazonas, Venezuela. Area-based R _d was positively related to area-based leaf N within functional groups and for all species pooled, but not when comparing among species within any site. At all sites, mass-based R _d (R _d-mass) decreased sharply with increasing leaf life-span and was positively related to SLA and mass-based A _max and leaf N (leaf N _mass). These intra-biome relationships were similar in shape and slope among sites, where in each case we compared species belonging to different plant functional groups. Significant R _d-mass−N _mass relationships were observed in all functional groups (pooled across sites), but the relationships differed, with higher R _d at any given leaf N in functional groups (such as forbs) with higher SLA and shorter leaf life-span. Regardless of biome or functional group, R _d-mass was well predicted by all combinations of leaf life-span, N _mass and/or SLA (r ²≥ 0.79, P < 0.0001). At any given SLA, R _d-mass rises with increasing N _mass and/or decreasing leaf life-span; and at any level of N _mass, R _d-mass rises with increasing SLA and/or decreasing leaf life-span. The relationships between R _d and leaf traits observed in this study support the idea of a global set of predictable interrelationships between key leaf morphological, chemical and metabolic traits. Received: 23 May 1997 / Accepted: 16 December 1997     

Leaf-trait relationships of <Emphasis Type="Italic">Quercus liaotungensis</Emphasis> along an altitudinal gradient in Dongling Mountain,Beijing

Few studies have examined leaf-trait relationships in the distribution of individual species along an environmental gradient. Here we address the issue by testing for the leaf-trait relationships of Quercus liaotungensis, a dominant deciduous woody species in northern China, along an altitudinal gradient in Dongling Mountain, Beijing. These leaf traits included specific leaf area (SLA), leaf dry matter content (LDMC), and leaf nitrogen, phosphorus, and potassium concentration on mass basis (N_mass, P_mass and K_mass, respectively). Along the altitudinal gradient, negative relationships between SLA and LDMC and N_mass were found, and N_mass, P_mass and K_mass correlated with each other positively. Relationship between N_mass and P_mass was stronger than the ones between N_mass and K_mass, and between P_mass and K_mass. The weak and negative relationship between SLA and N_mass might result from trade-offs that limit photosynthesis and water use efficiency along the altitudinal gradient, suggesting many environmental factors of local site being the collective forcing of drivers. Out of our expectations, N_mass and P_mass related very weakly to soil N and P, and no significant relationship between K_mass and soil K was found along elevation. These relationships could be used to predict the productivity of the population with changing environment in this region.     

A paradox of leaf-trait convergence: why is leaf nitrogen concentration higher in species with higher photosynthetic capacity?

It is well known that leaf photosynthesis per unit dry mass (A_mass) is positively correlated with nitrogen concentration (N_mass) across naturally growing plants. In this article we show that this relationship is paradoxical because, if other traits are identical among species, plants with a higher A_mass should have a lower N_mass, because of dilution by the assimilated carbon. To find a factor to overcome the dilution effect, we analyze the N_mass–A_mass relationship using simple mathematical models and literature data. We propose two equations derived from plant-growth models. Model prediction is compared with the data set of leaf trait spectrum obtained on a global scale. The model predicts that plants with a higher A_mass should have a higher specific nitrogen absorption rate in roots (SAR), less biomass allocation to leaves, and/or greater nitrogen allocation to leaves. From the literature survey, SAR is suggested as the most likely factor. If SAR is the sole factor maintaining the positive relationship between N_mass and A_mass, the variation in SAR is predicted to be much greater than that in A_mass; given that A_mass varies 130-fold, SAR may vary more than 2000-fold. We predict that there is coordination between leaf and root activities among species on a global scale. Kouki Hikosaka is the recipient of the BSJ Award for Young Scientist, 2006.     

Leaf habit does not predict leaf functional traits in cerrado woody species

Plant species with a high leaf life span (LLS) commonly have a low specific leaf area (SLA), leaf nitrogen per unit mass (N), and phosphorous concentration (P), whereas species with low LLS have a high SLA, N and P. However, LLS tends to be longer in species growing in low-nutrient soils and, therefore, differences in LLS and other leaf traits may not be consistent with a plant classification according to leaf habit. Here we investigated whether leaf habit is consistent with leaf economic spectrum trade-offs in cerrado (a Neotropical savanna) woody species. We analyzed the SLA, N and P of 125 woody species with a distinct leaf habit (deciduous, semideciduous, brevideciduous or evergreen). We also gathered data on the LLS (33 species), maximum net photosynthesis per leaf area (A_area, 56 species) and per leaf mass (A_mass, 31 species), comprising the most extensive database analyzed so far for the cerrado. Differences among leaf habit groups were tested using generalized linear mixed models and ANOVA. We did not find differences in SLA and N among species with a distinct leaf habit, but deciduous species had a higher leaf P concentration than evergreens. Species did not differ in LLS and A_mass, but A_area varied among groups. Semideciduous species had higher A_area values than deciduous and brevideciduous species, but all other groups had similar A_area values. Because of the small difference in the LLS, SLA, leaf N, leaf P and maximum net photosynthesis, we argue that deciduous, brevideciduous, semideciduous and evergreen species may not constitute different functional groups in cerrado woody species.     

Intraspecific variability in functional traits matters: case study of Scots pine

Although intraspecific trait variability is an important component of species ecological plasticity and niche breadth, its implications for community and functional ecology have not been thoroughly explored. We characterized the intraspecific functional trait variability of Scots pine (Pinus sylvestris) in Catalonia (NE Spain) in order to (1) compare it to the interspecific trait variability of trees in the same region, (2) explore the relationships among functional traits and the relationships between them and stand and climatic variables, and (3) study the role of functional trait variability as a determinant of radial growth. We considered five traits: wood density (WD), maximum tree height (H _max), leaf nitrogen content (N_mass), specific leaf area (SLA), and leaf biomass-to-sapwood area ratio (B _L:A _S). A unique dataset was obtained from the Ecological and Forest Inventory of Catalonia (IEFC), including data from 406 plots. Intraspecific trait variation was substantial for all traits, with coefficients of variation ranging between 8 % for WD and 24 % for B _L:A _S. In some cases, correlations among functional traits differed from those reported across species (e.g., H _max and WD were positively related, whereas SLA and N_mass were uncorrelated). Overall, our model accounted for 47 % of the spatial variability in Scots pine radial growth. Our study emphasizes the hierarchy of factors that determine intraspecific variations in functional traits in Scots pine and their strong association with spatial variability in radial growth. We claim that intraspecific trait variation is an important determinant of responses of plants to changes in climate and other environmental factors, and should be included in predictive models of vegetation dynamics.     

氮磷添加对金露梅叶片化学计量及光合特征的影响

通过三种养分添加处理,氮添加(5、10和15 g??m-2)、磷添加(梯度同氮添加)、氮磷同时添加[(5 g N＋5 g P)??m-2、(10 g N＋10 g P)??m-2、(15 g N＋15 g P)??m-2],对照(无养分添加),探讨养分添加对金露梅叶片性状氮含量(Nmas )、磷含量(Pmas )、氮磷比(N∶P)、比叶重(LMA)、净光合速率(Pn )和光合氮利用效率(PNUE)的影响,以及各性状之间的相互关系.结果表明：在处理水平上,除N或P显著提高金露梅叶片的N∶P外,氮、磷添加对叶片其它性状无显著影响;不同氮、磷处理下添加水平对金露梅叶片的Nmas、N∶P、Pn和PNUE均有显著影响,随着养分水平提高,各性状的变化模式各不相同,叶片Pmas无明显变化,而叶片LMA虽有降低的趋势但不显著.回归分析表明,叶片Pmas与Nmas之间呈显著正相关(R2＝0．347,P＜0．001),叶片Nmas 与N∶P之间也呈显著正相关(R2＝0．018,P＜0．05),而叶片Pmas与N∶P呈显著负相关(R2＝0．505,P＜0．001);叶片LMA与Pn之间显著负相关(R2＝0．02,P＜0．05),而与PNUE之间显著正相关(R2＝0．077,P＜0．001).这表明在一定范围内,环境变化可以改变金露梅叶片的养分保持能力、光合能力以及养分利用效率.     

Photosynthesis, leaf morphology and chemistry of Pinus koraiensis and Quercus mongolica in broadleaved Korean pine mixed forest

Leaf traits and physiology are species-specific and various with canopy position and leaf age. Leaf photosynthesis, morphology and chemistry in the upper and lower canopy positions of Pinus koraiensis Sieb. et Zucc and Quercus mongolica Fisch. ex Turoz in broadleaved Korean pine forest were determined in September 2009. Canopy position did not significantly affect light-saturated photosynthetic rate based on unit area (P _area) and unit dry mass (P _mass), apparent quantum yield (α), light compensation point (LCP), light saturation point (LSP); total nitrogen (N_m), phosphorus (P_m), carbon (C_m), and chlorophyll content (Chl_m) per unit dry mass; leaf dry mass per unit area (LMA) and photosynthetic nitrogen-use efficiency (PNUE) for P. koraiensis current-year needles and Q. mongolica leaves. While in P. koraiensis one-year-old needles, P _area, P _mass, α and LCP in the upper canopy were lower than those in the lower canopy. The needles of P. koraiensis had higher Cm and LMA than leaves of Q. mongolica, but P _mass, Chl_m and PNUE showed opposite trend. There were no differences in P _area, LSP, N_m, and P_m between the two species. Needle age significantly influenced photosynthetic parameters, chemistry and LMA of P. koraiensis needles except LCP, LSP and C_m. In contrast to LMA, P _area, P _mass, N_m, P_m, Chl_m, and PNUE of one-year-old needles were significantly lower than those of current-year needles for P. koraiensis. The negative correlations between LMA and P _mass, N_m, P_m, Chl_m, and positive correlations between P _mass and N_m, P_m, Chl_m were found for P. koraiensis current-year needles and Q. mongolica leaves. Our results indicate that leaf nitrogen and phosphorus contents and nutrient absorption from soil are similar for mature P. koraiensis and Q. mongolica growing in the same environment, while difference in carbon content between P. koraiensis and Q. mongolica may be attributed to inherent growth characteristics.     

唐古特白刺叶功能性状沿气候梯度的变异特征

叶功能性状与植物的生长对策及资源利用效率密切相关,研究叶功能性状沿气候梯度的变异特征能为理解植物对气候变化的响应机制提供一种简便可行的测定指标。以我国西北荒漠地区广泛分布的唐古特白刺（Nitraria tangutorum）为研究对象,对其比叶面积（SLA）、单位质量和单位面积叶氮含量（N_mass、N_area）、单位质量和单位面积叶建成成本（CC_mass、CC_area）进行测定,分析这些叶功能性状及性状相关关系沿气候梯度的变异特征。结果表明,唐古特白刺叶功能性状（CC_area除外）在气候梯度下存在显著差异,其中,温度是决定唐古特白刺SLA变化的主要因子,SLA随着温度的增加而增加;降水和温度对唐古特白刺N_mass、N_area和CC_mass均有显著影响,N_mass和N_area随着降水和温度的增加而降低,而CC_mass呈增加趋势。沿气候梯度,唐古特白刺SLA-N_mass、CC_mass-N_mass和CC_area-N_area的线性正相关关系发生平移,导致在相同SLA、CC_mass和CC_area下,降水和温度较低的地区具有更高的N_mass和N_area。这一结果表明唐古特白刺能通过调节叶功能性状之间的关系来适应气候的变化,并形成性状间的最佳功能组合。     

Photosynthesis-nitrogen relations in Amazonian tree species

The relationships between leaf nitrogen (N), specific leaf area (SLA) (an inverse index of leaf thickness or density), and photosynthetic capacity (A_max) were studied in 23 Amazonian tree species to characterize scaling in these properties among natural populations of leaves of different ages and light microenvironments, and to examine how variation within species in N and SLA can influence the expression of the A_max-to-N relationship on mass versus area bases. The slope of the A_max-N relationship, change in A per change in N (mol CO₂ gN^-1 s^-1), was consistently greater, by as much as 300%, when both measures were expressed on mass rather than area bases. The x-intercept of this relationship (N-compensation point) was generally positive on a mass but not an area basis. In this paper we address the causes and implications of such differences. Significant linear relationships (p<0.05) between mass-based leaf N (N_mass) and SLA were observed in 12 species and all 23 regressions had positive slopes. In 13 species, mass-based A_max (A_mass) was positively related (p<0.05) with SLA. These patterns reflect the concurrent decline in N_mass and SLA with increasing leaf age. Significant (p<0.05) relationships between area-based leaf N (N_area) and SLA were observed in 18 species. In this case, all relationships had negative slopes. Taken collectively, and consistent in all species, as SLA decreased (leaves become thicker) across increasing leaf age and light gradients, N_mass also decreased, but proportionally more slowly, such that N_area increased. Due to the linear dependence of A_mass on N_mass and a negative 4-intercept, thicker leaves (low SLA) therefore tend, on average, to have lower N_mass and A_mass but higher N_area than thinner leaves. This tendency towards decreasing A_mass with increasing N_area, resulting in a lower slope of the A_max-N relationship on an area than mass basis in 16 of 17 species where both were significant. For the sole species exception (higher area than mass-based slope) variation in N_area was related to variation in N_mass and not in SLA, and thus, these data are also consistent with this explanation. The relations between N, SLA and A_max explain how the rate of change in A_max per change in N can vary three-fold depending on whether a mass or area mode of expression is used.     

Productivity and thallus toughness trade‐off relationship in marine macroalgae from the Japan Sea

Trade‐offs are considered key to understanding mechanisms supporting the coexistence of multiple plant species. Thus, understanding the mechanisms underlying trade‐offs is expected to contribute to conservation and management of macroalgal beds composed of diverse macroalgae of rocky shore ecosystems. To test the occurrence of trade‐offs between productivity and thallus toughness as well as pair‐wise thallus trait relationships that are expected to indirectly relate to any trade‐offs, traits and relationships for 13 species of macroalgae from the central area along the Japan Sea coast of Honshu, Japan were examined. In each species we examined for photosynthetic capacity per unit biomass (as A_mass) and nitrogen (i.e., photosynthetic nitrogen‐use efficiency, PNUE), nitrogen content (as N_mass), thallus mass per unit thallus area (as TMA) and force required to penetrate the thallus (as F_p, a common index of leaf toughness in land plants by punch test). A significant negative correlation indicating a trade‐off between productivity and thallus toughness was found between A_mass or PNUE and F_p. Pair‐wise relationships that were expected to indirectly relate to the trade‐off were as follows. A_mass was positively correlated with N_mass. Thalli with high N_mass extensively utilizing nitrogen in the photosynthetic parts, and consequently exhibiting elevated metabolic rates. Moreover, thalli with high N_mass tended to be associated with low TMA, and N_mass decreased with increasing TMA. A significant negative correlation was observed between TMA and A_mass or PNUE because of the linkage of high A_mass or PNUE with high N_mass and high N_mass associated with low TMA, while a significant positive correlation was observed between TMA and F_p. The two correlations indicate a physiological and structural trade‐off, which underlies the interdependency of thallus traits. Results of multivariate analyses also indicated that the thallus traits interdependently vary across a single axis based on the trade‐off.     

Leaf attributes and tree growth in a tropical dry forest

Questions: How are leaf attributes and relative growth rate (RGR) of the dominant tree species of tropical deciduous forest (TDF) affected by seasonal changes in soil moisture content (SMC)? What is the relationship of functional attributes with each other? Can leaf attributes singly or in combination predict the growth rate of tree species of TDF? Location: Sonebhadra district of Uttar Pradesh, India. Methods: Eight leaf attributes, specific leaf area (SLA); leaf carbon concentration (LCC); leaf nitrogen concentration (LNC); leaf phosphorus concentration (LPC); chlorophyll concentration (Chl), mass‐based stomatal conductance (Gs_mass); mass based photosynthetic rate (A_mass); intrinsic water use efficiency (WUEi); and relative growth rate (RGR), of six dominant tree species of a dry tropical forest on four sites were analysed for species, site and season effects over a 2‐year period. Step‐wise multiple regression was performed for predicting RGR from mean values of SMC and leaf attributes. Path analysis was used to determine which leaf attributes influence RGR directly and which indirectly. Results: Species differed significantly in terms of all leaf attributes and RGR. The response of species varied across sites and seasons. The attributes were positively interrelated, except for WUEi, which was negatively related to all other attributes. The positive correlation was strongest between Gs_mass and A_mass and the negative correlation was strongest between Gs_mass and WUEi. Differences in RGR due to site were not significant when soil moisture was controlled, but differences due to season remained significant. The attributes showed plasticity across moisture gradients, which differed among attributes and species. Gs_mass was the most plastic attribute. Among the six species, Terminalia tomentosa exhibited the greatest plasticity in six functional attributes. In the step‐wise multiple regression, A_mass, SLA and Chl among leaf attributes and SMC among environmental factors influenced the RGR of tree species. Path analysis indicated the importance of SLA, LNC, Chl and A_mass in determining RGR. Conclusion: A _mass, SMC, SLA and Chl in combination can be used to predict RGR but could explain only three‐quarters of the variability in RGR, indicating that other traits/factors, not studied here, are also important in modulating growth of tropical trees. RGR of tree species in the dry tropical environment is determined by soil moisture, whereas the response of mature trees of different species is modulated by alterations in key functional attributes such as SLA, LNC and Chl.     

Leaf anatomical structures of <Emphasis Type="Italic">Paphiopedilum</Emphasis> and <Emphasis Type="Italic">Cypripedium</Emphasis> and their adaptive significance

Paphiopedilum and Cypripedium are closely related in phylogeny, but have contrasting leaf traits and habitats. To understand the divergence in leaf traits of Paphiopedilum and Cypripedium and their adaptive significance, we analyzed the leaf anatomical structures, leaf dry mass per area (LMA), leaf lifespan (LL), leaf nitrogen concentration (N _mass), leaf phosphorus concentration (P _mass), mass-based light-saturated photosynthetic rate (A _mass), water use efficiency (WUE), photosynthetic nitrogen use efficiency (PNUE) and leaf construction cost (CC) for six species. Compared with Cypripedium, Paphiopedilum was characterized by drought tolerance derived from its leaf anatomical structures, including fleshy leaves, thick surface cuticles, huge adaxial epidermis cells, lower total stoma area, and sunken stomata. The special leaf structures of Paphiopedilum were accompanied by longer LL; higher LMA, WUE, and CC; and lower N _mass, P _mass, A _mass, and PNUE compared with Cypripedium. Leaf traits in Paphiopedilum helped it adapt to arid and nutrient-poor karst habitats. However, the leaf traits of Cypripedium reflect adaptations to an environment characterized by rich soil, abundant soil water, and significant seasonal fluctuations in temperature and precipitation. The present results contribute to our understanding of the divergent adaptation of leaf traits in slipper orchids, which is beneficial for the conservation of endangered orchids.     

入侵种银胶菊和三叶鬼针草与本地种气体交换特性的比较

以菊科入侵植物银胶菊(Parthenium hysterophorus)和三叶鬼针草(Bidens pilosa)以及与其共生的菊科本地植物小蓟(Cirsium setosum)为对象,比较了3种植物气体交换参数和叶片特性的差异。结果表明,银胶菊和三叶鬼针草的净光合速率(net photosynthetic rate,P_n)、叶绿素含量、比叶面积(specific leaf area,SLA)、叶片单位质量P含量(leaf P content per unit mass,P_(mass))、光合能量利用效率(photosynthetic energy use efficiency,PEUE)和光合氮利用效率(photosynthetic nitrogen use efficiency,PNUE)均显著高于小蓟。植物叶片P_n与水分利用效率(water use efficiency,WUE)、叶片P_(mass)、SLA呈极显著正相关,植物叶片单位质量N含量(leaf P content per unit mass,N_(mass))与叶片SLA、单位质量建成成本(leaf construction cost per unit mass,CC_(mass))、叶绿素含量呈极显著正相关。与本地植物相比,较高的气体交换参数和叶片生化指标有可能是银胶菊和三叶鬼针草成功入侵的原因之一。     

不同种源五角枫幼苗叶片N、P化学计量学特征对氮添加的响应

不同土壤氮环境下植物对氮沉降的响应趋势不同,因此研究不同种源地的五角枫(Acer mono)对氮沉降的响应机理具有重要的意义。为了深入了解不同种源五角枫对氮沉降的适应和响应机制,主要通过3个种源的五角枫幼苗氮添加控制实验,分析不同种源的五角枫幼苗比叶面积(SLA)和叶片干物质含量(LMDC)以及叶片N、P含量和N∶P对氮添加的响应规律。结果表明:(1)不同氮添加处理下,内蒙古种源五角枫幼苗(NW)和山西种源五角枫幼苗(SW)的LDMC均显著低于对照,NW和SW在N4条件下表现为显著的促进作用。随着氮添加浓度的增加,NW与北京种源的五角枫幼苗(BW)的SLA在N4条件下显著增加;SW的SLA在N2的水平下显著增加。(2)氮添加下,只有SW的P_(mass)在N1水平下显著增加,N_(mass)在N1和N3水平下表现为显著增加。说明在低氮水平下,能够促进SW对P的吸收,从而增加植物体内对P的积累。同时在低氮和中高氮条件下,能够促进SW对N的吸收,说明SW能较好的适应低氮和高氮环境,使其生存能力更强。(3)分析变异来源及各指标的相关性得出,氮添加条件下,五角枫的SLA、LDMC、P_(area)及N∶P与氮添加处理呈显著的相关关系,而SLA、LDMC、N_(mass)、P_(mass)、Narea、P_(area)和N∶P对氮添加处理和种源的交互作用均未表现出相关关系,说明引起五角枫各性状主要受氮添加处理影响。(4)综合各指标的变化,氮添加条件下,NW的LDMC显著降低,SLA显著增大,同时叶N含量、叶P含量以及N∶P比较高;SW的LDMC显著降低,SLA增大,同时叶N含量、叶P含量表现为显著增大,结合叶经济谱理论分析得出,SW和NW属于"快速投资-收益"型,而BW倾向于"缓慢投资-收益"型。因此,不同种源的五角枫对于氮沉降的响应存在差异,并且持续的氮添加已经改变了五角枫的适应性。