张掖湿地芦苇比叶面积和水分利用效率的关系

密度制约下植物比叶面积与水分利用效率的关系,对于认识土壤-植物-大气的物质循环和能量流动机制具有重要意义。采用样方调查法,研究了3种密度(高密度Ⅰ:210—230株/m~2;中密度Ⅱ:130—150株/m~2;低密度Ⅲ:50—70株/m~2)条件下芦苇种群比叶面积(SLA)和水分利用效率(WUE)的关系。结果表明:随着芦苇种群密度的逐渐降低,湿地群落的土壤含水量逐渐减小,芦苇的株高、叶面积、叶干重、SLA和蒸腾速率(Tr)均呈逐渐减小的趋势,净光合速率(Pn)、叶厚度和WUE呈逐渐增加的趋势;不同密度条件下湿地植物芦苇比叶面积(SLA)与水分利用效率(WUE)的关系存在显著差异(P0.05),在高密度(Ⅰ)与低密度(Ⅲ)样地,芦苇SLA与WUE呈极显著负相关关系(P0.01);在中密度(Ⅱ)样地,二者呈显著负相关关系(P0.05)。不同种群密度的湿地生境中,芦苇通过调整叶片形态构造,使比叶面积和水分利用效率形成了相反的变化趋势,反映了植物适应光照条件、土壤含水量等异质性环境因子的资源利用策略和光合产物积累模式。     

叶面积指数的研究和应用进展

对叶面积指数(LAI)提出50多年来,在植物光合作用、蒸腾作用、联系光合和蒸腾的关系和构成生产力基础的研究,在林分、景观以及地区尺度上对碳、能量、水分通量的研究,借助遥感技术建立森林生态系统的生长模型以及研究森林生态系统的能量和水分交换等方面的研究和应用进展进行了综述。LAI作为进行植物群体和群落生长分析的一个重要参数,已在农业、果树业、林业以及生物学、生态学等领域得到广泛应用。     

秦王川盐沼湿地芦苇叶片比叶面积与光合效率的关联分析

植物比叶面积(SLA)与光合效率的关联性分析,有助于理解植物叶片的光合产物分配与能量分配之间的权衡机制。该研究以秦王川国家湿地公园芦苇(Phragmites australis)种群为研究对象,沿芦苇群落末端至水域边缘分别依次设置:(I:土壤电导率(EC) 2.3—2.8 ms/cm)、(II:1.8—2.2 ms/cm)、(III:0.8—1.5 ms/cm) 3个试验样地,研究了秦王川盐沼湿地不同土壤盐分条件下芦苇叶片SLA与光合效率的关系。结果表明:随着土壤盐分含量的降低,湿地群落的高度、地上生物量逐渐增大,密度、光合有效辐射(PAR)呈逐渐减小的趋势;叶面积、株高、净光合速率(P_n)、水分利用效率(WUE)和蒸腾速率(T_r)均呈逐渐增大的趋势,叶厚度逐渐减小,比叶面积呈先增加后减少的变化趋势,叶干重的变化趋势与比叶面积完全相反;芦苇叶片实际光合效率(Y(II))、光化学淬灭系数(Q_P)和电子传递速率(ETR)呈先减少后增大的变化趋势,非光化学猝灭系数(NPQ)、调节性能量耗散的量子产额(Y(NPQ))和最大光化学效率(F_v/F_m)呈先增加后减少的变化趋势,非调节性能量耗散的量子产量(Y(NO))呈逐渐增大的变化趋势。在3个样地中,芦苇SLA与Y(II)和ETR之间均呈极显著负相关关系(P0.01),SMA斜率的绝对值均呈先增大后减少的变化趋势。随土壤含盐量的梯度性变化,芦苇种群适时调整叶片构件模式以改变叶片电子传递速率和实际光化学效率,实现植物光合效率的最大化,反映了盐沼湿地植物的特殊生存策略和叶片构件模式。     

不同CO2浓度变化下干旱对冬小麦叶面积指数的影响差异

叶面积指数是作物生长状况的一个重要表征参数,也是研究陆地生态系统的一个重要的参数.当今世界温室气体排放逐年上升,气候变暖趋势明显,对气候变化敏感的农业将受到影响.在全球变化的背景下,采用农业技术转移决策支持系统(DSSAT)系统,通过在黄淮海平原典型站点模拟3种CO2浓度条件下冬小麦在水分充足和水分亏缺2种情境下的生长过程,分析不同CO2浓度下水分亏缺对冬小麦叶面积指数的影响差异.研究发现,CO2浓度升高对叶面积指数增长有促进作用,且在干旱情况下对叶面积指数的正效应比湿润情况下更为明显,在CO2浓度倍增条件下,发生水分亏缺的作物叶面积指数数倍增长.研究结论有助于分析CO2浓度变化对农作物生长过程的影响,为农田水分管理提供依据,又为估算叶面积指数提出了一种模型的方法.     

模拟喀斯特生境条件下干旱胁迫对青冈栎苗木的影响

为了研究喀斯特"双层"地质结构对植物的影响,以及在干旱环境下,喀斯特地区植物的生理变化与适应策略,建立"土壤层-岩石(石灰岩)层-岩溶水层"水分供应分层模拟柱,对‘土壤层’设置不同水分梯度,种植青冈栎(Cyclobalanopsis glauca)苗木进行干旱胁迫试验,测定青冈栎的叶面积、比叶面积、生物量、渗透调节物质含量、相对叶绿素含量。结果表明:在模拟柱‘岩溶水层’加水条件下,植物根系能下扎至‘岩溶水层’,‘土壤层’干旱胁迫对青冈栎的生理变化没有产生影响;在‘岩溶水层’无水条件下,青冈栎生理变化受‘土壤层’干旱胁迫影响显著,‘土壤层’水分含量越低,青冈栎的叶面积、生物量、叶绿素含量、叶片相对含水量越小,青冈栎干鲜比、根冠比、可溶性糖、丙二醛和脯氨酸含量越高。青冈栎幼苗利用岩溶水层水分是适应喀斯特干旱环境的重要策略。     

西藏紫花针茅叶功能性状沿降水梯度的变化

植物叶功能性状与环境因子的关系是近10年来植物生态学的研究热点。该文以广泛分布于青藏高原干旱、半干旱草地的优势植物种紫花针茅(Stipa purpurea)为研究对象, 沿降水梯度(69-479 mm)系统测定了日土、改则、珠峰、当雄和纳木错5个调查地点紫花针茅比叶面积(SLA)、单位重量和单位面积叶氮含量(N_mass, N_area)、叶密度和厚度等叶功能性状以及土壤全氮含量等因子, 试图验证干旱胁迫地区同一物种内SLA-N_mass关系沿降水梯度的策略位移现象是否具有普遍性, 并对是否出现策略位移现象提出可能的解释。研究结果表明: 1) SLA和N_mass与生长季温度和降水以及土壤全氮含量均没有显著关系, SLA与N_mass的关系在干旱半干旱区(年降水/蒸发比< 0.11)与半湿润区(年降水/蒸发比> 0.11)之间并没有出现典型的位移现象; 2)叶密度是决定半湿润区SLA变化的主导因子, 而叶厚度则是干旱半干旱区SLA变化的控制因子, 两者与SLA均呈负相关, 随着温度增加或降水减少, 叶厚度增加而叶密度降低, 导致SLA随温度和降水变化不明显; 3)半湿润区的叶密度增加引起N_area增加, 而干旱半干旱区的叶厚度增加并没有造成N_area的显著变化, 导致N_area沿降水梯度没有显著变化; 4)紫花针茅地上生物量与N_area具有显著正相关关系, 表明N_area的增加有助于提高植被生产力。结果表明, 在干旱胁迫下, 植物通过增加叶厚度来维持不变的N_area可能有助于保持与较湿润地区相似的光合生产和水分利用效率。叶厚度和叶密度对比叶面积的相对影响在干旱半干旱区与半湿润区之间发生转变, 这为进一步检测高寒草地植被的水分限制阈值提供了新思路。     

高寒退化草地狼毒种群个体大小与茎、叶的异速生长

叶片和茎的生长规律决定植物对光截取的模式以及与外界环境的物质交换面积,其生物量分配是研究植物生活史对策的一个重要途径.在石羊河上游高寒退化草地,采用标准化主轴估计(standardized major axis estimation,SMA)方法,研究了狼毒(Stellera chamaejasme)种群茎大小对茎与叶生物量分配的影响,以及对叶面积支持效率的影响.结果表明:狼毒种群茎干重与比叶重(LMA)、单叶面积、叶干重呈异速生长关系,SMA斜率分别为0.781(95％的置信区间,CI=O.71 ～0.94)、0.824(CI=0.77 ～0.95)和0.856(CI=0.79 ～0.99).狼毒种群中较大茎的狼毒植株具有较高的叶生物量分配比例和叶面积支持效率,同时具有较高的比叶重,说明较大的狼毒采取以提高资源利用效率为主的适应策略,而相对较小的狼毒采取以快速生长为主的适应策略.     

半干旱黄土丘陵区人工林叶面积特征

 该文通过对黄土丘陵区4个密度的刺槐（Robinia pseudoacacia）人工林、3个密度的侧柏（Platycladus orientalis）人工林生长季叶面积变化的研究,揭示了不同密度林分叶面积生长与林分密度的关系;通过对灌木生长季叶面积变化的研究,建立了灌木柠条(Caragana korshinskii)、沙棘（Hippophae rhamnoides）和紫穗槐（Amorpha fruticosa）叶面积与叶鲜重、枝条基径的经验公式,为半干旱区灌木生长调查提供了一种方便、快捷的方法。结果表明：1）刺槐和侧柏各密度林分的单株林木叶面积和叶面积指数均在9月达到最大值,其中刺槐林叶面积指数峰值可达到10.5,侧柏峰值可达到3.2;灌木柠条、沙棘和紫穗槐叶面积和叶面积指数都在8月份达到各自的最大值,柠条、沙棘和紫穗槐的叶面积指数峰值分别为1.1 95、1.123和1.882;2）刺槐叶面积与叶鲜重具有极显著相关的幂函数关系,侧柏、柠条、沙棘、紫穗槐叶面积与叶鲜重具有极显著相关的线性函数关系,其中柠条枝条基径与叶面积还具有极显著相关的幂函数关系,沙棘、紫穗槐枝条基径与叶面积还具有极显著相关的线性函数关系;3）黄土丘陵区,由于林地土壤水分条件的限制,承载力有限。人工林进入生长盛期后, 不同密度刺槐和侧柏林分叶面积指数趋于一致,与最初的造林密度和现存密度没有关系。在不同密度的刺槐和侧柏林分间,单株叶面积与其林分密度成反比。在对上述结果分析的基础上得出：黄土丘陵区,由于林地土壤水分条件的限制,承载力有限。该文所研究的刺槐和侧柏各林分均已达到了当地土地承载力的上限,基于提高单株林木质量的考虑,建议刺槐林郁闭后的密度不超过833株&;#8226;hm^-2,侧柏则不超过1 111株&;#8226;hm^－2。如以全林分生物量为目标,林分密度也可适当减小。     

张掖湿地芨芨草叶大小和叶脉密度的权衡关系

叶大小-叶脉密度的权衡关系是植物叶经济谱理论的基础, 对理解资源竞争条件下植物叶片的物理构建与生理代谢的关系具有重要的意义。该文采用标准化主轴估计(standardized major axis estimation, SMA)的方法, 按芨芨草(Achnatherum splendens)株丛密度设置I (>12丛·m^-2)、II (8-12丛·m^-2)、III (4-8丛·m^-2)和IV (<4丛·m^-2) 4个密度梯度, 以叶面积和叶干质量分别表示叶大小, 对张掖洪泛平原湿地不同密度条件下芨芨草种群的叶大小和叶脉密度的关系进行研究。结果表明: 随着芨芨草株丛密度的降低, 湿地群落的土壤含水量逐渐减小、土壤电导率逐渐增加, 芨芨草的净光合速率(P_n)、蒸腾速率(T_r)和分枝数呈先增大后减小的趋势, 叶面积、叶干质量、比叶面积和株高呈逐渐减小趋势、光合有效辐射(PAR)和叶脉密度呈逐渐增加趋势; 芨芨草叶大小和叶脉密度在高密度(I)和低密度(IV)样地均呈极显著负相关关系(p < 0.01), 中密度(II、III)样地二者呈显著负相关关系(p < 0.05); 叶大小和叶脉密度回归方程的SMA斜率在不同密度样地均显著小于-1 (p < 0.05), 即芨芨草叶大小和叶脉密度呈“此消彼长”的权衡关系。在高密度湿地群落芨芨草倾向于大叶片低叶脉密度的叶片构建模式, 在低密度湿地群落选择小叶片高叶脉密度的异速生长模式, 体现了密度制约下湿地植物的生物量分配格局和资源利用对策。     

不同密度条件下芨芨草空间格局对环境胁迫的响应

基于小尺度上植物间相互作用与空间格局的高度相关性,选用可避免环境异质性影响的K2点格局函数,研究了沙枣-芨芨草群落中63个芨芨草样方(3密度条件×3生境条件×7重复)中芨芨草种群在小尺度下(0—0.5 m)的空间格局,探讨了在3种密度条件(高、中、低)及3种生境条件下(冠盖区、过渡区、空旷区),芨芨草种群空间格局对土壤理化性质胁迫(盐分、有机质及容重)程度的响应。研究结果表明,就土壤盐分,养分含量及土壤容重而言,土壤理化性质胁迫程度沿冠盖区、过渡区和空旷区增加的趋势。相应地,芨芨草种群呈聚集分布的样方比例在沿冠盖区(6/21)较低,而在过渡区(11/21)和空旷区(11/21)较高。然而,在不同密度条件下,芨芨草种群空间格局对土壤性质胁迫的响应不同。在低密度条件下,芨芨草种群在冠盖区多数为聚集分布(4/7),在过渡区和空旷区全部为聚集分布;中密度条件下,芨芨草聚集分布样方比例沿冠盖区-过渡区-空旷区梯度增加(分别为2/7,3/7,4/7),但低于同等胁迫条件下低密度样方中聚集分布数量;高密度条件下,在各胁迫条件下,除了一个过渡区高密度样方,其余芨芨草种群均为随机分布。总体上,随着环境胁迫增强,在中低密度下,芨芨草种群趋向于聚集分布;但在高密度下,芨芨草种群均以随机分布为主。此外,芨芨草种群空间格局随密度变化趋势比随土壤理化性质胁迫梯度变化趋势更加明显,可能表明相对于土壤理化性质胁迫程度,芨芨草种群密度对其空间格局影响更大。因此,在考虑芨芨草种群空间格局对环境胁迫的响应时,应当考虑种群密度因素。     

Comparison of CO<Subscript>2</Subscript> and H<Subscript>2</Subscript>O fluxes over grassland vegetations measured by the eddy-covariance technique and by open system chamber

Measurements of CO₂ and H₂O fluxes were carried out using two different techniques—eddy-covariance (EC) and open system gas exchange chamber (OC)—during two-years’ period (2003–2004) at three different grassland sites. OC measurements were made during fourteen measurement campaigns. We found good agreement between the OC and EC CO₂ flux values (n = 63, r ² = 0.5323, OC F_CO2 = −0.6408+0.9508 EC F_CO2). The OC F_H2O values were consistently lower than those measured by the EC technique, probably caused by the air stream difference inside and outside the chamber. Adjusting flow rate within the chamber to the natural conditions would be necessary in future OC measurements. In comparison with EC, the OC proved to be a good tool for gas exchange measurements in grassland ecosystems.     

The scaling of leaf area and mass: the cost of light interception increases with leaf size

For leaves, the light-capturing surface area per unit dry mass investment (specific leaf area, SLA) is a key trait from physiological, ecological and biophysical perspectives. To address whether SLA declines with leaf size, as hypothesized due to increasing costs of support in larger leaves, we compiled data on intraspecific variation in leaf dry mass (LM) and leaf surface area (LA) for 6334 leaves of 157 species. We used the power function LM=alpha LAbeta to test whether, within each species, large leaves deploy less surface area per unit dry mass than small leaves. Comparing scaling exponents (beta) showed that more species had a statistically significant decrease in SLA as leaf size increased (61) than the opposite (7) and the average beta was significantly greater than 1 (betamean=1.10, 95% CI 1.08-1.13). However, scaling exponents varied markedly from the few species that decreased to the many that increased SLA disproportionately fast as leaf size increased. This variation was unrelated to growth form, ecosystem of origin or climate. The average within-species tendency found here (allometric decrease of SLA with leaf size, averaging 13%) is in accord with concurrent findings on global-scale trends among species, although the substantial scatter around the central tendency suggests that the leaf size dependency does not obligately shape SLA. Nonetheless, the generally greater mass per unit leaf area of larger than smaller leaves directly translates into a greater cost to build and maintain a unit of leaf area, which, all else being equal, should constrain the maximum leaf size displayed.     

Temporal dynamics and spatial variability in the enhancement of canopy leaf area under elevated atmospheric CO2

Increased canopy leaf area (L) may lead to higher forest productivity and alter processes such as species dynamics and ecosystem mass and energy fluxes. Few CO₂ enrichment studies have been conducted in closed canopy forests and none have shown a sustained enhancement of L. We reconstructed 8 years (1996–2003) of L at Duke's Free Air CO₂ Enrichment experiment to determine the effects of elevated atmospheric CO₂ concentration ([CO₂]) on L before and after canopy closure in a pine forest with a hardwood component, focusing on interactions with temporal variation in water availability and spatial variation in nitrogen (N) supply. The dynamics of L were reconstructed using data on leaf litterfall mass and specific leaf area for hardwoods, and needle litterfall mass and specific leaf area combined with needle elongation rates, and fascicle and shoot counts for pines. The dynamics of pine L production and senescence were unaffected by elevated [CO₂], although L senescence for hardwoods was slowed. Elevated [CO₂] enhanced pine L and the total canopy L (combined pine and hardwood species; P<0.050); on average, enhancement following canopy closure was ～16% and 14% respectively. However, variation in pine L and its response to elevated [CO₂] was not random. Each year pine L under ambient and elevated [CO₂] was spatially correlated to the variability in site nitrogen availability (e.g. r²=0.94 and 0.87 in 2001, when L was highest before declining due to droughts and storms), with the [CO₂]‐induced enhancement increasing with N (P=0.061). Incorporating data on N beyond the range of native fertility, achieved through N fertilization, indicated that pine L had reached the site maximum under elevated [CO₂] where native N was highest. Thus closed canopy pine forests may be able to increase leaf area under elevated [CO₂] in moderate fertility sites, but are unable to respond to [CO₂] in both infertile sites (insufficient resources) and sites having high levels of fertility (maximum utilization of resources). The total canopy L, representing the combined L of pine and hardwood species, was constant across the N gradient under both ambient and elevated [CO₂], generating a constant enhancement of canopy L. Thus, in mixed species stands, L of canopy hardwoods which developed on lower fertility sites (～3 g N inputs m^?2 yr^?1) may be sufficiently enhanced under elevated [CO₂] to compensate for the lack of response in pine L, and generate an appreciable response of total canopy L (～14%).     

Size dependence of leaf area and the mass of component organs during a course of self-thinning in a hinoki (Chamaecyparis obtusa) seedling population

The allometric equation, y=gx^h, was applied monthly to the relationships between two different dimensions of tree seedlings of hinoki cypress (Chamaecyparis obtusa) during a course of self-thinning from April 1990 to March 1991 to detect differences in biomass allocation among individuals. As the h-value in the allometry of crown length and seedling height was greater than unity for all seasons, crown ratio became greater as seedling height increased. Leaf weight ratio increased with increasing seedling size because the h-value in the allometry of leaf dry weight and whole seedling dry weight was greater than unity in every month. Therefore, smaller seedlings are disadvantageous to photosynthetic production by leaves. In contrast, the leaf area ratio was constant irrespective of seedling size because the h-value in the allometry of leaf area and whole seedling dry weight was nearly equal to unity in most seasons. In addition, because the h-value in the allometry of leaf area and leaf dry weight was less than unity in all seasons, specific leaf area decreased with an increase in leaf dry weight, indicating that smaller seedlings adapt to low light environments by possessing shade leaves. Root weight ratio decreased with increasing seedling size because the h-value in the allometry of root dry weight and whole seedling dry weight was less than unity in most seasons.     

从异龄叶性状分析几种常绿阔叶植物变异与关联

该研究从个体与物种两个水平分析福建省鬼洞山中亚热带常绿阔叶次生林中七种阔叶树种异龄叶的平均叶面积(MLA)、比叶面积(SLA)和叶干物质含量(LDMC)三个性状的变异及关联,探讨叶性状物质分配策略对植物生活策略优化的限制性影响。结果表明:(1)对七种阔叶树种的个体和物种水平变异系数而言,均为MLA(82.9%;76.9%)SLA(38.9%;35.5%)LDMC(25.4%;23.8%);在个体和物种水平上当年生叶片MLA变异系数(71.5%;64.0%)小于往年生叶片(72.2%;65.8%),SLA、LDMC值则相反。(2)在个体和物种水平,当年生叶MLA对往年生叶MLA变异的解释率分别为50.1%和61.5%,当年生叶SLA对往年生叶SLA变异的解释率分别为56.6%和77.0%,当年生叶LDMC对往年生叶LDMC变异的解释率分别为51.7%和68.3%。(3)7种亚热带常绿阔叶植物异龄叶MLA、SLA与LDMC变化规律说明,当年生叶与往年生叶投资相同干物质,当年叶可形成更大的叶面积,且叶面积建成的消耗较往年叶小。研究认为,异龄叶性状在异龄叶间存在变异与关联,叶面积形成过程中生物量建成与消耗的协调可能影响植物叶片的发育。     

A Comparison of Photosynthesis in Endangered and Non-Endangered Plants Changium smyrnioides and Anthriscus sylvestris

Changium smyrnioides Wolff. and Anthriscus sylvestris (L.) Hoffm. have similar photosynthetic characters; they use radiant energy in winter and early spring effectively, but cannot take full advantage of higher irradiance after spring. The specific leaf area (SLA), leaf area ratio (LAR), and leaf mass ratio (LMR) of C. smyrnioides were lower than those of A. sylvestris. The photosynthetic period of C smyrnioides was about 160 d shorter than that of A. sylvestris, causing the total photosynthetic production of C. smyrnioides to be lower than that of A. sylvestris. Hence if C. smyrnioides is disturbed, it could not recover within a short period.     

Scaling‐up from leaf to canopy‐aggregate properties in sclerophyll shrub species

Abstract: Plant species vary widely in their average leaf lifespan (LL) and specific leaf area (SLA, leaf area per dry mass). The negative LL–SLA relationship commonly seen among species represents an important evolutionary trade‐off, with higher SLA indicating greater potential for fast growth (higher rate of return on a given investment), but longer LL indicating a longer duration of the revenue stream from that investment. We investigated how these leaf‐economic traits related to aggregate properties of the plant crown. Across 14 Australian sclerophyll shrub species, those with long LL accumulated more leaf mass and leaf area per unit ground area. Light attenuation through their canopies was more severe. Leaf accumulation and light attenuation were more weakly related to SLA than to LL. The greater accumulation of foliage in species with longer LL and lower SLA may counterbalance their generally lower photosynthetic rates and light‐capture areas per gram of leaf.     

Scale‐dependent trait correlations in a temperate tree community

Recent investigations of relationships among plant traits have generated important insights into plant form and function. However, relationships involving leaf area, leaf shape and plant height remain poorly resolved. Previous work has also focused on correlations between average trait values for individual species. It is unclear whether similar relationships occur within species. We searched for novel plant trait correlations by comparing leaf area, leaf circularity, specific leaf area (SLA) and plant height among 16 common woody plant species from a temperate forest in New Zealand. Analyses were conducted both within species (intra‐specifically) and among species (inter‐specifically) to determine whether trait correlations were scale dependent. Leaf area was unrelated to other leaf traits inter‐specifically. However, leaf area declined with plant height and increased with SLA intra‐specifically. Leaf circularity decreased with plant height inter‐specifically, but increased with plant height intra‐specifically. SLA increased with plant height both inter‐ and intra‐specifically. Leaf circularity increased with SLA inter‐specifically, but decreased with SLA intra‐specifically. Overall results showed that leaf shape, SLA and plant height are interrelated. However, intra‐specific relationships often differed substantially from inter‐specific relationships, suggesting that the processes shaping relationships between this suite of plant traits are scale‐dependent.     

Adaptive survival mechanisms and growth limitations of small-stature herb species across a plant diversity gradient

Several biodiversity experiments have shown positive effects of species richness on aboveground biomass production, but highly variable responses of individual species. The well-known fact that the competitive ability of plant species depends on size differences among species, raises the question of effects of community species richness on small-stature subordinate species. We used experimental grasslands differing in species richness (1-60 species) and functional group richness (one to four functional groups) to study biodiversity effects on biomass production and ecophysiological traits of five small-stature herbs (Bellis perennis, Plantago media, Glechoma hederacea, Ranunculus repens and Veronica chamaedrys). We found that ecophysiological adaptations, known as typical shade-tolerance strategies, played an important role with increasing species richness and in relation to a decrease in transmitted light. Specific leaf area and leaf area ratio increased, while area-based leaf nitrogen decreased with increasing community species richness. Community species richness did not affect daily leaf carbohydrate turnover of V. chamaedrys and P. media indicating that these species maintained efficiency of photosynthesis even in low-light environments. This suggests an important possible mechanism of complementarity in such grasslands, whereby smaller species contribute to a better overall efficiency of light use. Nevertheless, these species rarely contributed a large proportion to community biomass production or achieved higher yields in mixtures than expected from monocultures. It seems likely that the allocation to aboveground plant organs to optimise carbon assimilation limited the investment in belowground organs to acquire nutrients and thus hindered these species from increasing their performance in multi-species mixtures.