Large variation in whole-plant water-use efficiency among tropical tree species

It is well known that whole-plant water-use efficiency (transpiration efficiency of carbon gain, TE(C)) varies among plant species with different photosynthetic pathways. However, less is known of such variation among tree species within the C(3) group. Here we measured the TE(C) of seven C(3) tropical tree species. Isotopic analyses (delta(13)C, delta(18)O, and delta(15)N) and elemental analyses (carbon and nitrogen) were undertaken to provide insight into sources of variation in TE(C). Plants were grown over several months in approx. 80% full sunlight in individual 38-l containers in the Republic of Panama. Soil moisture content was nonlimiting. Significant variation was observed in TE(C) among the C(3) tree species. Values ranged from 1.6 mmol C mol(-1) H(2)O for teak (Tectona grandis) to 4.0 mmol C mol(-1) H(2)O for a legume, Platymiscium pinnatum. Variation in TE(C) was correlated with both leaf N concentration, a proxy for photosynthetic capacity, and oxygen-isotope enrichment, a proxy for stomatal conductance. The TE(C) varied with C-isotope discrimination within species, but the relationship broke down among species, reflecting the existence of species-specific offsets.     

Conifers, angiosperm trees, and lianas: growth, whole-plant water and nitrogen use efficiency, and stable isotope composition ({delta}13C and {delta}18O) of seedlings grown in a tropical environment

Seedlings of several species of gymnosperm trees, angiosperm trees, and angiosperm lianas were grown under tropical field conditions in the Republic of Panama; physiological processes controlling plant C and water fluxes were assessed across this functionally diverse range of species. Relative growth rate, r, was primarily controlled by the ratio of leaf area to plant mass, of which specific leaf area was a key component. Instantaneous photosynthesis, when expressed on a leaf-mass basis, explained 69% of variation in r (P < 0.0001, n = 94). Mean r of angiosperms was significantly higher than that of the gymnosperms; within angiosperms, mean r of lianas was higher than that of trees. Whole-plant nitrogen use efficiency was also significantly higher in angiosperm than in gymnosperm species, and was primarily controlled by the rate of photosynthesis for a given amount of leaf nitrogen. Whole-plant water use efficiency, TE(c), varied significantly among species, and was primarily controlled by c(i)/c(a), the ratio of intercellular to ambient CO(2) partial pressures during photosynthesis. Instantaneous measurements of c(i)/c(a) explained 51% of variation in TE(c) (P < 0.0001, n = 94). Whole-plant (13)C discrimination also varied significantly as a function of c(i)/c(a) (R(2) = 0.57, P < 0.0001, n = 94), and was, accordingly, a good predictor of TE(c). The (18)O enrichment of stem dry matter was primarily controlled by the predicted (18)O enrichment of evaporative sites within leaves (R(2) = 0.61, P < 0.0001, n = 94), with some residual variation explained by mean transpiration rate. Measurements of carbon and oxygen stable isotope ratios could provide a useful means of parameterizing physiological models of tropical forest trees.     

Physiological and isotopic (δ13C and δ18O) responses of three tropical tree species to water and nutrient availability

Water-use efficiency and stable isotope composition were studied in three tropical tree species. Seedlings of Tectona grandis , Swietenia macrophylla and Platymiscium pinnatum were grown at either high or low water supply, and with or without added fertilizer. These three species previously exhibited low, intermediate and high whole-plant water-use efficiency ( TE ) when grown at high water supply in unfertilized soil. Responses of TE to water and nutrient availability varied among species. The TE was calculated as experiment-long dry matter production divided by cumulative water use. Species-specific offsets were observed in relationships between TE and whole-plant ¹³C discrimination (Δ¹³C_p). These offsets could be attributed to a breakdown in the relationship between Δ¹³C_p and the ratio of intercellular to ambient CO₂ partial pressures ( c _i/ c _a) in P. pinnatum , and to variation among species in the leaf-to-air vapour pressure difference ( v ). Thus, a plot of v · TE against c _i/ c _a showed a general relationship among species. Relationships between δ ¹⁸O of stem dry matter and stomatal conductance ranged from strongly negative for S. macrophylla to no relationship for T. grandis . Results suggest inter-specific variation among tropical tree species in relationships between stable isotope ratios ( δ ¹³C and δ ¹⁸O) and the gas exchange processes thought to affect them.     

δ<Superscript>13</Superscript>C and water-use efficiency in Australian grasstrees and South African conifers over the last century

Annual or biannual time courses of plant delta13C (delta13C(p)) over the last century (70-100 years) were recorded for leafbases of four grasstrees (Xanthorrhoea preissii) at four sites in mediterranean Australia and wood of four conifers (Widdringtonia cedarbergensis) at two sites in mediterranean South Africa. There was a strong downward trend of 2-5.5(per thousand ) from 1935 to 1940 to the present in the eight plants. Trends were more variable from 1900 to 1940 with plants at two sites of each species showing an upward trend of 1-2.5 per thousand. Accepting that delta13C of the air (delta13C(a)) fell by almost 2 per thousand over the last century, the ratio of leaf intercellular CO2 to atmospheric CO2 (c(i)/c(a)) rose in five plants and remained unchanged in three over that period. Changes in c(i)/c(a) rather than delta13C(a) were more closely correlated with changes in delta13C(p) and accounted for 6.7-71.8% (22.6 c(i)/c(a)) and 28.2-93.3% (delta13C(a)) of the variation in delta13C(p). We doubt that possible changing patterns of rainfall, water availability, temperature, shade, air pollution or clearing for agriculture have contributed to the overall trend for c(i)/c(a) to rise over time. Instead, we provide evidence (concentrations of Fe and Mn in the grasstree leafbases) that decreasing photosynthetic capacity associated with falling nutrient availability due to the reduced occurrence of fire may have contributed to rising c(i)/c(a). Intrinsic water-use efficiency (W(i)) as a function of (c(a)-c(i)) usually increased linearly over the period, with the two exceptions explained by their marked increase in c(i)/c(a). We conclude that grasstrees may provide equivalent delta13C(p )and W(i) data to long-lived conifers and that their interpretation requires a consideration of the causes of variation in both c(i)/c(a )and delta13C(a).     

Discrimination in the dark. Resolving the interplay between metabolic and physical constraints to phosphoenolpyruvate carboxylase activity during the crassulacean acid metabolism cycle

A model defining carbon isotope discrimination (delta13C) for crassulacean acid metabolism (CAM) plants was experimentally validated using Kalanchoe daigremontiana. Simultaneous measurements of gas exchange and instantaneous CO2 discrimination (for 13C and 18O) were made from late photoperiod (phase IV of CAM), throughout the dark period (phase I), and into the light (phase II). Measurements of CO2 response curves throughout the dark period revealed changing phosphoenolpyruvate carboxylase (PEPC) capacity. These systematic changes in PEPC capacity were tracked by net CO2 uptake, stomatal conductance, and online delta13C signal; all declined at the start of the dark period, then increased to a maximum 2 h before dawn. Measurements of delta13C were higher than predicted from the ratio of intercellular to external CO2 (p(i)/p(a)) and fractionation associated with CO2 hydration and PEPC carboxylations alone, such that the dark period mesophyll conductance, g(i), was 0.044 mol m(-2) s(-1) bar(-1). A higher estimate of g(i) (0.085 mol m(-2) s(-1) bar(-1)) was needed to account for the modeled and measured delta18O discrimination throughout the dark period. The differences in estimates of g(i) from the two isotope measurements, and an offset of -5.5 per thousand between the 18O content of source and transpired water, suggest spatial variations in either CO2 diffusion path length and/or carbonic anhydrase activity, either within individual cells or across a succulent leaf. Our measurements support the model predictions to show that internal CO2 diffusion limitations within CAM leaves increase delta13C discrimination during nighttime CO2 fixation while reducing delta13C during phase IV. When evaluating the phylogenetic distribution of CAM, carbon isotope composition will reflect these diffusive limitations as well as relative contributions from C3 and C4 biochemistry.     

Determinants of Leaf Litter Nutrient Cycling in a Tropical Rain Forest: Soil Fertility Versus Topography

We investigated the influence of landscape-level variation in soil fertility and topographic position on leaf litter nutrient dynamics in a tropical rain forest in Costa Rica. We sampled across the three main edaphic conditions (ultisol slope, ultisol plateau, and inceptisol) to determine the effect of soil nutrients on leaf litter nutrient concentrations while controlling for topography, and to examine topographic effects while controlling for soil nutrients. Both leaf litter macronutrient [phosphorus (P), nitrogen (N), sulfur (S), calcium (Ca), potassium (K), magnesium (Mg)] and micronutrient concentrations were quantified throughout a 4-year period. Leaf litter [P], [N] and [K] varied significantly among soil types. The variation in [P], [N], and [K] was explained by soil fertility alone. Leaf litter [S], [Ca], and [Mg] did not vary among the three soil types. Macronutrient (P, K, Mg, S, Ca) concentrations in the leaf litter were much less variable than those of Fe and Al. Lower variability in essential plant nutrients suggests a great deal of plant control over the amount of nutrients resorbed before senescense. Leaf litter macronutrient concentrations varied significantly over the 4-year period, but the temporal variation did not differ among the three edaphic types as anticipated. Hence, although the magnitude of nutrient fluxes may be controlled by local factors such as soil fertility, temporal patterns are likely regulated by a common environmental variable such as precipitation or temperature.     

Leaf chlorophyll concentration relates to transpiration efficiency in peanut

Two pot experiments were conducted in two different seasons at the University of Agricultural Science, Bangalore, India, to study (a) the relationship between chlorophyll concentration (by measuring the leaf light‐transmittance characteristics using a SPAD metre) and transpiration efficiency (TE) and (b) the effect of leaf N on chlorophyll and TE relationship in peanut. In Experiment (Expt) I, six peanut genotypes with wide genetic variation for the specific leaf area (SLA) were used. In Expt II, three non‐nodulating isogenic lines were used to study the effect of N levels on leaf chlorophyll concentration–TE relationship without potential confounding effects in biological nitrogen fixation. Leaf N was manipulated by applying N fertiliser in Expt II. Chlorophyll concentration, TE (g dry matter kg^?1 of H₂O transpired, measured using gravimetric method), specific leaf nitrogen (g N m^?2, SLN), SLA (cm² g^?1), carbon isotope composition (Δ¹³C) were determined in the leaves sampled during the treatment period (35–55 days after sowing) in the two experiments. Results showed that the leaf chlorophyll concentration expressed as soil plant analytical development (SPAD) chlorophyll metre reading (SCMR) varied significantly among genotypes in Expt I and as a result of N application in Expt II. Changes in leaf N levels were strongly associated with changes in SCMR, TE and Δ¹³C. In both the experiments, a significant positive relationship between SCMR and TE with similar slopes but differing intercepts was noticed. However, correction of TE for seasonal differences in vapour pressure deficit (VPD) between the two experiments resulted in a single and stronger relationship between SCMR and TE. There was a significant inverse relationship between SCMR and Δ¹³C, suggesting a close linkage between chlorophyll concentration and Δ¹³C in peanut. This study provides the first evidence for a significant positive relationship between TE and leaf chlorophyll concentration in peanut. The study also describes the effect of growing environment on the relationships among SLA, SLN and SCMR.     

Temporal variation in δ<Superscript>13</Superscript>C of ecosystem respiration in the Pacific Northwest: links to moisture stress

We measured seasonal and interannual variations in delta(13)C values within the carbon reservoirs (leaves and soil) and CO(2) fluxes (soil and ecosystem respired CO(2)) of an old growth coniferous forest in the Pacific Northwest USA with relation to local meteorological conditions. There were significant intra-annual and interannual differences in the carbon isotope ratios of CO(2) respired at both the ecosystem (delta(13)C(R)) and the soil levels (delta(13)C(R-soil)), but only limited variations in the carbon isotope ratios of carbon stocks. The delta(13)C(R) values varied by as much as 4.4 per thousand over a growing season, while delta(13)C(R-soil )values changed as much as 6.2 per thousand. The delta(13)C of soil organic carbon (delta(13)C(SOC)) and needle organic carbon (delta(13)C(P)) exhibited little or no significant changes over the course of this study. Carbon isotope discrimination within leaves (Delta(p)) showed systematic decreases with increased canopy height, but remained fairly constant throughout the year (Delta(p)=17.9 per thousand -19.2 per thousand at the top of the canopy, Delta(p)=19.6 per thousand -20.9 per thousand at mid-canopy, Delta(p)=23.3 per thousand -25.1 per thousand at the canopy base). The temporal variation in the delta(13)C of soil and ecosystem respired CO(2) was correlated ( r=0.93, P<0.001) with soil moisture levels, with dry summer months having the most (13)C-enriched values. The dynamic seasonal changes in delta(13)C of respired CO(2) are hypothesized to be the result of fast cycling of recently fixed carbon back to the atmosphere. One scaling consequence of the seasonal and interannual variations in delta(13)C(R) is that inversion-based carbon-cycle models dependent on observed atmospheric CO(2) concentration and isotope values may be improved by incorporating dynamic delta(13)C(R) values to interpret regional carbon sink strength.     

Leaf nonstructural carbohydrate concentrations of understory woody species regulated by soil phosphorus availability in a tropical forest

Leaf soluble sugars and starch are important components of nonstructural carbohydrates (NSCs), which are crucial for plant growth, development, and reproduction. Although there is a large body of research focusing on the regulation of plant NSC (soluble sugars and starch) concentrations, the response of foliar NSC concentrations to continuous nitrogen (N) and phosphorus (P) addition is still unclear, especially in tropical forests. Here, we used a long‐term manipulative field experiment to investigate the response of leaf NSC concentrations to continuous N and P addition (3‐, 5‐, and 8‐year fertilization) in a tropical forest in southern China. We found significant species‐specific variation in leaf NSC concentrations in this tropical forest. Phosphorus addition dramatically decreased both leaf soluble sugar and starch concentrations, while N addition had no significant effects on leaf soluble sugar and starch concentrations. These results suggest that, in plants growing in P‐limiting tropical soil, leaf NSC concentrations are regulated by soil P availability rather than N availability. Moreover, the negative relationships between NSC concentrations and leaf mass per area (LMA) revealed that NSCs could supply excess carbon (C) for leaf expansion under P addition. This was further supported by the increased structural P fraction after P fertilization in our previous study at the same site. We conclude that soil P availability strongly regulates leaf starch and soluble sugar concentrations in the tropical tree species included in this study. The response of leaf NSC concentrations to long‐term N and P addition can reflect the close relationships between plant C dynamics and soil nutrient availability in tropical forests. Maintaining relatively higher leaf NSC concentrations in tropical plants can be a potential mechanism for adapting to P‐deficient conditions.     

集约化生产对农田土壤碳氮含量及δ13C和δ15N同位素丰度的影响

集约化生产下农田土壤碳、氮含量变化是衡量土壤肥力持久性的重要指标.对常规水稻-蚕豆轮作地、露地蔬菜地、3年塑料大棚地和10年以上塑料大棚地的土壤pH、电导率(EC)、土壤有机碳(SOC)和总氮(TN)含量及δ13C和δ15N同位素丰度进行测定,研究了集约化生产程度对土壤特性的影响.结果表明:与水稻-蚕豆轮作地相比,露地蔬菜地、3年塑料大棚地和10年以上塑料大棚地0 ～20 cm耕层土壤pH分别降低1.1、0.8和0.7,而土壤EC分别是水稻-蚕豆轮作地的4.2、4.9和5.2倍;土壤碳、氮含量随塑料大棚地生产年限的增加总体上呈先增大后减小的趋势.与水稻-蚕豆轮作地相比,10年以上塑料大棚地0～20、20～40、40 ～60、60 ～ 80、80 ～ 100 cm土层的土壤SOC含量分别下降了54％、46％、60％、63％和59％,土壤TN含量分别下降了53％、53％、71％、82％和85％.农田集约化生产程度显著影响土壤SOC、TN含量和δ13C、δ15N丰度,土壤δ13C丰度与SOC含量呈显著负相关.土壤δ13C丰度可作为评价农田土壤碳循环受人为干扰强度的指标.     

Non-omnivorous generality promotes population stability

Using a combination of stable isotope analysis of delta13C and delta15N and long-term census data on population abundances for meiofauna in tropical aquatic rock pools, we provide evidence that species which exhibit greater variation in delta13C, an indication of a greater range of distinct carbon sources in their diet, have more stable populations than species with lower variation in delta13C. This link between increased isotope variability and reduced population variability, however, did not hold for delta15N. This suggests that increases in population stability were due to non-omnivorous feeding on multiple carbon sources within a trophic level rather than omnivorous feeding on multiple carbon sources across trophic levels. Our findings corroborate MacArthur's original hypothesis that populations that can access a greater range of resources are more stable than those which consume a more restricted range of resources.     

Does the Prostrate-leaved Geophyte Brunsvigia orientalis Utilize Soil-derived CO2 for Photosynthesis?

BACKGROUND AND AIMS: A test was made of the hypothesis that the prostrate growth habit of the leaves of the geophyte Brunsvigia orientalis enables utilization of soil-derived CO(2) and is related to the presence of lysigenous air-filled channels characteristic of B. orientalis leaves. METHODS: Brunsvigia orientalis was sampled at a field site. Leaf anatomy, stomatal density, leaf/soil gas exchange characteristics and soil atmosphere and leaf delta(13)C isotope abundances were examined. KEY RESULTS: The leaves of B. orientalis have large lysigenous air-filled channels separating the upper and lower surfaces of the leaves. The upper surface comprised approx. 70 % of the leaf mass and 75 % of the leaf N (mmol g(-1)). Between 20 % and 30 % of the stomatal conductance and CO(2) assimilation was through the lower surface of the leaf. CO(2) efflux rates from the soil surface were up to 5.4 micromol m(-2) s(-1) while photosynthetic fluxes through the lower surface of the leaves were approx. 7 micromol m(-2) s(-1). However, the utilization of soil-derived CO(2) only altered the leaf delta(13)C isotope abundance of the prostrate leaves by a small amount. Using delta(13)C values it was estimated that 7 % of the leaf tissue C was derived from soil-derived CO(2). CONCLUSIONS: A small proportion of photosynthetically fixed CO(2) was derived from the soil, with minimal associated transpirational H(2)O loss into the space between the leaf and soil. The soil-derived CO(2), taken up through the lower surface was probably assimilated by the palisade tissue in the upper surface of the leaf which was exposed to sunlight and where most of the leaf N was located. The occurrence of lysigenous air channels in the leaves may provide longitudinal strength without impaired transfer of CO(2) taken up through the lower surface to the upper surface.     

Leaf functional response to increasing atmospheric CO(2) concentrations over the last century in two northern Amazonian tree species: a historical δ(13) C and δ(18) O approach using herbarium samples

We assessed the extent of recent environmental changes on leaf morphological (stomatal density, stomatal surface, leaf mass per unit area) and physiological traits (carbon isotope composition, δ(13)C(leaf) , and discrimination, Δ(13)C(leaf) , oxygen isotope composition, δ(18)O(leaf) ) of two tropical rainforest species (Dicorynia guianensis; Humiria balsamifera) that are abundant in the Guiana shield (Northern Amazonia). Leaf samples were collected in different international herbariums to cover a 200 year time-period (1790-2004) and the whole Guiana shield. Using models describing carbon and oxygen isotope fractionations during photosynthesis, different scenarios of change in intercellular CO(2) concentrations inside the leaf (C(i)), stomatal conductance (g), and photosynthesis (A) were tested in order to understand leaf physiological response to increasing air CO(2) concentrations (C(a)). Our results confirmed that both species displayed physiological response to changing C(a) . For both species, we observed a decrease of about 1.7‰ in δ(13)C(leaf) since 1950, without significant change in Δ(13)C(leaf) and leaf morphological traits. Furthermore, there was no clear change in δ(18)O(leaf) for Humiria over this period. Our simulation approach revealed that an increase in A, rather than a decrease in g, explained the observed trends for these tropical rainforest species, allowing them to maintain a constant ratio of C(i)/C(a) .     

海拔梯度对长白山北坡岳桦水分利用效率的影响

以分布于长白山北坡海拔1800～2050 m的岳桦林为对象,以叶片碳稳定同位素(δ13C)值作为岳桦长期水分利用效率指示值,探讨海拔梯度对岳桦林水分利用效率的影响.结果表明:随海拔升高,岳桦林土壤体积含水量、比叶质量极显著增加,而叶片含水量和土壤温度显著降低;岳桦叶片δ13C值与海拔呈极显著正相关,增幅为1.013‰·(100 m)-1,与土壤体积含水量、比叶质量呈显著正相关,与生长季土壤平均温度、叶片含水量呈显著负相关.温度不是长白山林线的唯一限制性因子,海拔梯度上水热条件的差异及其交互作用可能造成岳桦生长期间的生理干旱.     

凋落物化学组成对土壤微生物学性状及土壤酶活性的影响

通过模拟试验的方法研究了单一施加杉木(Cunn inghan ia lancceola ta(L am b)Hook.)叶凋落物,杉木(C.lancceola ta)和桤木(A lnus crem astogyne Burk ill)混合凋落物,杉木(C.lancceola ta)和枫香(L iqu id am ba f orm osana H ance)混合凋落物,杉木(C.lancceola ta)、桤木(A.crem astogyne)、枫香(L.f orm osana)混合凋落物对土壤化学性状和土壤微生物量碳、代谢熵(qCO2)、土壤酶活性的影响。研究结果表明,土壤微生物学性状比土壤化学性状对不同凋落物处理的效应反应更敏感;与单一杉木叶凋落物比较,混合凋落物处理的土壤微生物量碳明显增加,土壤脲酶、蔗糖酶、脱氢酶活性升高;土壤代谢熵(qCO2)和土壤多酚氧化酶活性有下降趋势;另外,研究结果也表明,不同树种的叶凋落物混合对土壤质量的影响存在差异,有桤木叶的混合凋落物对土壤质量的改善效果似乎更明显。     

Responses of legume versus nonlegume tropical tree seedlings to elevated CO2 concentration

We investigated responses of growth, leaf gas exchange, carbon-isotope discrimination, and whole-plant water-use efficiency (W(P)) to elevated CO(2) concentration ([CO(2)]) in seedlings of five leguminous and five nonleguminous tropical tree species. Plants were grown at CO(2) partial pressures of 40 and 70 Pa. As a group, legumes did not differ from nonlegumes in growth response to elevated [CO(2)]. The mean ratio of final plant dry mass at elevated to ambient [CO(2)] (M(E)/M(A)) was 1.32 and 1.24 for legumes and nonlegumes, respectively. However, there was large variation in M(E)/M(A) among legume species (0.92-2.35), whereas nonlegumes varied much less (1.21-1.29). Variation among legume species in M(E)/M(A) was closely correlated with their capacity for nodule formation, as expressed by nodule mass ratio, the dry mass of nodules for a given plant dry mass. W(P) increased markedly in response to elevated [CO(2)] in all species. The ratio of intercellular to ambient CO(2) partial pressures during photosynthesis remained approximately constant at ambient and elevated [CO(2)], as did carbon isotope discrimination, suggesting that W(P) should increase proportionally for a given increase in atmospheric [CO(2)]. These results suggest that tree legumes with a strong capacity for nodule formation could have a competitive advantage in tropical forests as atmospheric [CO(2)] rises and that the water-use efficiency of tropical tree species will increase under elevated [CO(2)].     

Effects of intraleaf variations in carbonic anhydrase activity and gas exchange on leaf C18OO isoflux in Zea mays

Variation in the C18OO content of atmospheric CO2 (delta18Oa) can be used to distinguish photosynthesis from soil respiration, which is based on carbonic anhydrase (CA)-catalyzed 18O exchange between CO2 and 18O-enriched leaf water (delta18Ow). Here we tested the hypothesis that mean leaf delta18Ow and assimilation rates can be used to estimate whole-leaf C18OO flux (isoflux), ignoring intraleaf variations in CA activity and gas exchange parameters. We observed variations in CA activity along the leaf (> 30% decline from the leaf center toward the leaf ends), which were only partially correlated to those in delta18Ow (7 to 21 per thousand), delta18O and delta13C of leaf organic matter (25 to 30 per thousand and -12.8 to -13.2 per thousand, respectively), and substomatal CO2 concentrations (intercellular CO2 concentrations, c(i), at the leaf center were approximately 40% of those at the leaf tip). The combined effect of these variations produced a leaf-integrated isoflux that was different from that predicted based on bulk leaf values. However, because of canceling effects among the influencing parameters, isoflux overestimations were only approximately 10%. Conversely, use of measured parameters from a leaf segment could produce large errors in predicting leaf-integrated C18OO fluxes.     

Nutrient return with leaf litter fall in Fagus sylvatica forests across a soil fertility gradient

Leaf litter fall is an important nutrient flux in temperature deciduous forests which supplies a large part of the rapidly mineralisable nutrient fraction to the soil. This study investigates nutrient return with leaf litter fall in 36 old-growth forest stands of Fagus sylvatica across a broad gradient of soil fertility covering 9 mesozoic and kaenozoic parent material types (three limestones, two sandstones, two clay stones, one sand and one loess substrate). Study objectives were to analyse (i) the dependency of leaf litter nutrient concentrations on soil fertility, and (ii) the relationship between soil fertility and nutrient return with leaf litter at the stand level. Beech stands on the nine parent material types produced similar annual leaf litter masses irrespective of soil fertility or acidity. Leaf litter from the nine parent materials showed only minor variation with respect to N and K concentrations (factors of 1.5 and 1.4), moderate variation for Ca, Mg and P concentrations (factors of 2.2 to 2.9), and high variation for Al and Mn concentrations (factors of 6.7 and 10.5). Consequently, annual nutrient return with litter fall (leaf litter mass x litter nutrient concentration) was more similar among the parent materials for N (165–273 mmol m⁻² yr1) ⁻¹ and K (16–30 mm m⁻² yr⁻¹) than for Ca, P, Mg, Mn and Al. A possible explanation is increased N deposition in recent time. According to a correlation analysis, return rates of N, P, K and Mg (but not Ca) were independent of the pool size of the respective nutrient in the soil. N return rate was neither influenced by the soil pools of Nt, plant- available P (Pa) or exchangeable Ca, K and Mg, nor by soil acidity or the exchangeable Al pool. P return, in contrast, showed a negative relation to soil fertility. We hypothesize that nutrient fluxes with leaf litter fall do not necessarily reduce the fitness of tree populations as has been postulated from a tree-centred view. Rather, we suggest that nutrient fluxes with litter fall can increase, instead of decrease, plant fitness by improving nutrient availability in the densely rooted topsoil which reduces the roots’ carbon and nutrient costs of nutrient acquisition.     

Organic matter inputs shift soil enzyme activity and allocation patterns in a wet tropical forest

Soil extracellular enzymes mediate organic matter turnover and nutrient cycling yet remain little studied in one of Earth’s most rapidly changing, productive biomes: tropical forests. Using a long-term leaf litter and throughfall manipulation, we explored relationships between organic matter (OM) inputs, soil chemical properties and enzyme activities in a lowland tropical forest. We assayed six hydrolytic soil enzymes responsible for liberating carbon (C), nitrogen (N) and phosphorus (P), calculated enzyme activities and ratios in control plots versus treatments, and related these to soil biogeochemical variables. While leaf litter addition and removal tended to increase and decrease enzyme activities per gram soil, respectively, shifts in enzyme allocation patterns implied changes in relative nutrient constraints with altered OM inputs. Enzyme activity ratios in control plots suggested strong belowground P constraints; this was exacerbated when litter inputs were curtailed. Conversely, with double litter inputs, increased enzymatic investment in N acquisition indicated elevated N demand. Across all treatments, total soil C correlated more strongly with enzyme activities than soluble C fluxes, and enzyme ratios were sensitive to resource stoichiometry (soil C:N) and N availability (net N mineralization). Despite high annual precipitation in this site (MAP ~5 m), soil moisture positively correlated with five of six enzymes. Our results suggest resource availability regulates tropical soil enzyme activities, soil moisture plays an additional role even in very wet forests, and relative investment in C, N and P degrading enzymes in tropical soils will often be distinct from higher latitude ecosystems yet is sensitive to OM inputs.     

Stable isotopes as indicators of altitudinal distributions and movements in an Ecuadorean hummingbird community

Altitudinal migration and dispersal is an important component of the life history of several temperate and tropical birds but remains poorly understood due to the limited success of mark and recapture techniques. Stable isotopes of hydrogen (deltaD) in rainfall, and to a lesser extent, carbon (delta13C) in plants are known to change with altitude and hence may provide the basis of a technique for tracking the altitudinal movements in birds and other wildlife. We investigated the potential for this technique by measuring delta13C, deltaD, and delta15N values in tail feathers of eight species of hummingbirds ( Phaethornis malaris, P. syrmatophorus, P. guy, Adelomyia melanogenys, Coeligena torquata, C. lutetiae, Metallura baroni, M. williami) along an altitudinal gradient (300-3,290 m asl) in the Andes Mountains of Ecuador. Feather delta13C and deltaD values were correlated and each changed significantly with elevation above 400 m. In general, we found good agreement between feather deltaD values and those predicted from a generalized relationship of precipitation and surface water deltaD with altitude. Similarly, feather delta13C values showed an enrichment of approximately 1.5 per thousand per 1,000 m over the linear portion of the elevational response. Stable-nitrogen isotope values were variable, and so did not provide useful information on elevation in birds, apart from trophic effects. Overall there appears to be good potential for using the (deltaD, delta13C) stable isotope approach to track altitudinal movements and to elucidate previously unrecognized patterns of life history variation in both temperate and tropical species that migrate across elevational isotopic gradients.