Nutrients are assimilated efficiently by Lymantria dispar caterpillars from the mature leaves of trees in the Salicaceae

The efficient aquisition of nutrients from leaves by insect herbivores increases their nutrient assimilation rates and overall fitness. Caterpillars of the gypsy moth (Lymantria dispar L.) have high protein assimilation efficiencies (PAE) from the immature leaves of trees such as red oak (Quercus rubra) and sugar maple (Acer saccharum) (71–81%) but significantly lower PAE from their mature leaves (45–52%). By contrast to this pattern, both PAE and carbohydrate assimilation efficiencies (CAE) remain high for L. dispar larvae on the mature leaves of poplar (Populus alba × Populus tremula) grown in greenhouse conditions. The present study tests two alternative hypotheses: (i) outdoor environmental stresses cause decreased nutrient assimilation efficiencies from mature poplar leaves and (ii) nutrients in the mature leaves of trees in the poplar family (Salicaceae) remain readily available for L. dispar larvae. When poplar trees are grown in ambient outdoor conditions, PAE and CAE remain high (approximately 75% and 78%, respectively) in L. dispar larvae, in contrast to the first hypothesis. When larvae feed on the mature leaves of species in the Salicaceae [aspen (Populus tremuloides), cottonwood (Populus deltoides), willow (Salix nigra) and poplar], PAE and CAE also remain high (68–76% and 72–92%, respectively), consistent with the second hypothesis. Larval growth rates are strongly associated with protein assimilation rates, and more strongly associated with protein assimilation rates than with carbohydrate assimilation rates. It is concluded that tree species in the Salicaceae are relatively high‐quality host plants for L. dispar larvae, in part, because nutrients in their mature leaves remain readily available.     

Physiological factors affecting the rapid decrease in protein assimilation efficiency by a caterpillar on newly‐mature tree leaves

Lymantria dispar L. caterpillars have a decreased ability to assimilate protein from mature leaves of red oak (Quercus rubra) compared with young, expanding leaves. The present study determines whether the drop in protein assimilation efficiency (PAE) occurs during the rapid phase of leaf maturation. Several mechanisms that might account for decreased PAE are also examined: mature leaf tissues could resist being chewed efficiently, protein in mature leaf tissues could become difficult to extract, and other nutrients in mature leaves might become growth limiting. The entire seasonal decrease in PAE occurs rapidly (in less than 2 weeks), when the leaves finished expanding. The maturation process is characterized by increased levels of fibre and decreased levels of water but no significant changes in the levels of protein or carbohydrates. Despite increased fibre in mature leaves, they are not chewed into larger food particles than are immature leaves. Carbohydrate assimilation efficiencies remain high on mature leaves, and signs of limiting water levels in larvae of L. dispar on mature leaves are not observed. The most important finding in the present study is the decreased extractability of protein in food particles from mature leaves, which plays a major role in explaining the rapid decrease in PAE. It is hypothesized that structural changes in cell walls during the rapid process of leaf maturation decrease protein extractability, which, in turn, greatly decreases the nutritional quality of mature oak leaves for caterpillars. The results of the present study therefore suggest a general mechanism to help explain the widely documented decrease in the nutritional quality of the mature leaves of many tree species for herbivorous insects.     

Effects of leaf maturity and wind stress on the nutrition of the generalist caterpillar Lymantria dispar feeding on poplar

The growth rates of insect herbivores commonly decrease when they feed on mature leaves due to the combined effects of several nutritional and physiological mechanisms. Environmental stresses during leaf development may also decrease herbivore performance. The present study tests two main hypotheses to help clarify the importance of these factors for the nutrition and growth of an insect herbivore: (i) decreases in nutrient levels, consumption rates and nutrient assimilation efficiencies impact negatively on herbivores feeding on mature leaves and (ii) wind stress has a negative impact on herbivores feeding on mature leaves. The results show that mature poplar (Populus alba × Populus tremula) leaves have decreased levels of protein and increased levels of fibre, and that growth rates of gypsy moth (Lymantria dispar L.) are decreased on mature leaves in association with decreased consumption rates. However, in contrast to the first hypothesis, protein and carbohydrate are assimilated efficiently (74–82% and 84–87%, respectively) from immature and mature poplar leaves. The larvae are able to chew mature leaves as efficiently as immature leaves, potentially maximizing nutrient extraction. By contrast to the second hypothesis, wind‐stressed leaves have no significant detrimental effects on nutrient assimilation efficiencies, and the lower growth rates of L. dispar larvae feeding on mature wind‐stressed leaves can be explained by lower consumption rates. Therefore, the availability of nutrients to herbivores feeding on mature tree leaves is not necessarily impacted by lower assimilation efficiencies, even when leaves develop under wind stress. These results help explain some of the large variation between the nutritional qualities of trees for forest Lepidoptera.     

Phenology of nutritional differences between new and mature leaves and its effect on caterpillar growth

Abstract. 1. We determined the phenology of the shrub Spiraea latifolia Ait. Bork. (Rosaceae), which has indeterminate shoot growth, and the effects of phenological changes in leaf quality on growth rate of the early-spring feeding buckmoth caterpillars, Hemileuca lucina Hy. Edw. (Saturniidae).
2. Leaves, regardless of whether they were newly expanded or several weeks old, were tougher later in the growth season (mid-June) than similarly aged leaves collected earlier; correspondingly, water and nitrogen content for leaves of all ages declined through the larval period. By July, newly expanded leaves had no more nitrogen than mature leaves.
3. Relative growth rate of third instar larvae fed new leaves or a mixture of new and mature leaves in early June was higher than that of those fed mature leaves, and efficiency of conversion of digested food to biomass was higher for larvae fed new leaves than for those fed mature leaves or a mixture.
4. In another experiment, larvae were reared on new leaves through the fourth instar and then fed a diet of new, mature or a combination of new and mature leaves, a regimen that was similar to the phenologies of both plants and caterpillars in the field. There was no difference in time to pupation or pupal weights among these treatments.     

Effects of elevated temperature and carbon dioxide on the nutritional quality of leaves of oak (Quercus robur L.) as food for the Winter Moth (Operophtera brumata L.)

1. Pedunculate Oak trees were grown in ambient and elevated temperatures and CO₂. Leaves were fed to Winter Moth caterpillars reared either in constant conditions or with the trees (caged or on-tree).
2. Caterpillars in constant conditions ate the same mass and produced the same mass of faeces whether fed elevated or ambient temperature leaves. However, less was assimilated from elevated leaves, resulting in lighter pupae and fewer, lighter eggs.
3. Caterpillars in constant conditions ate more and produced more faeces when fed elevated CO₂ leaves than when fed ambient CO₂ leaves, but the mass assimilated and pupal mass were unchanged.
4. Caged caterpillars reared with the trees from which they were fed had constant pupal mass in all treatments, but pupated earlier at elevated temperature. Pupal mass was also unaffected when caterpillars fed on the trees.
5. Nitrogen was reduced in both elevated temperature and elevated CO₂ leaves. Increased fibre in the former prevented increased consumption and resulted in reduced pupal mass and fecundity. Reduced fibre in the latter allowed increased consumption, resulting in pupae of normal mass.
6. Despite the clear effect of nutrient quality, experiments rearing caterpillars and trees together suggest that anticipated climatic change will have no nutritional effect on Winter Moth development.     

A comparison of foliar nutrient concentration in trees from monsoon rainforest and savanna in northern Australia

Abstract Analysis of foliar nutrient concentrations revealed that two mesophyllic monsoon rainforest trees had higher concentrations of most nutrients in leaves than eight savanna species. One of the tested monsoon rainforest species with sclerophyllous leaves had similar nutrient concentrations to the savanna tree species. There were positive or no significant correlations between live foliar nutrient concentration and the percentage of nutrients withdrawn prior to litterfall. The nutrient concentration of litter was similar for most nutrients among tree species of monsoon rainforest or savanna. The results of this study suggest that the relative fertility of surface soils of monsoon rainforest compared with savanna is not determined by contrasting nutrient cycling strategies whereby monsoon rainforest tree species enrich their soils with relatively nutrient rich leaf litter relative to savanna tree species.     

Ontogenetic variation in digestion by the herbivorous lizard Ctenosaura pectinata

I tested the hypothesis that an animal with an ontogenetic diet shift must have different digestive efficiencies for foods that correspond to its diet shift, so that nutrient and energy extraction are maximized. The iguanine lizard Ctenosaura pectinata undergoes an ontogenetic diet shift from eating insects as a juvenile to plants as an adult. When fed six different pure foods from the natural diets of different age classes, C. pectinata assimilated nutrients and energy differently depending on food type and age class. Extraction of energy and nutrients in insect larvae was maximized by juvenile lizards. Calcium, phosphorus, and energy were readily assimilated from flowers and fruit by immature and adult lizards. Magnesium levels were highest in leaves and were extracted by immature and adult lizards, but xenobiotic effects of one plant leaf (Croton suberosus), eaten by adults, killed juvenile lizards. Although juvenile C. pectinata ate some flowers (Senna wislizenii) naturally, they were less efficient at digesting cell walls from these plant parts than were older lizards. Ontogenetic changes in ctenosaur digestive physiology were not the result of a trade-off involving ecological costs of different foods; rather, each age class preferred a diet that maximized its physiological benefit.     

Foliar demand and resource economy of nutrients in dry tropical forest species

Important phenological activities in seasonally dry tropical forest species occur within the hot‐dry period when soil water is limiting, while the subsequent wet period is utilized for carbon accumulation. Leaf emergence and leaf area expansion in most of these tree species precedes the rainy season when the weather is very dry and hot and the soil cannot support nutrient uptake by the plants. The nutrient requirement for leaf expansion during the dry summer period, however, is substantial in these species. We tested the hypothesis that the nutrients withdrawn from the senescing leaves support the emergence and expansion of leaves in dry tropical woody species to a significant extent. We examined the leaf traits (with parameters such as leaf life span, leaf nutrient content and retranslocation of nutrients during senescence) in eight selected tree species in northern India. The concentrations of N, P and K declined in the senescing foliage while those of Na and Ca increased. Time series observations on foliar nutrients indicated a substantial amount of nutrient resorption before senescence and a ‘tight nutrient budgeting’. The resorbed N‐mass could potentially support 50 to 100% and 46 to 80% of the leaf growth in terms of area and weight, respectively, across the eight species studied. Corresponding values for P were 29 to 100% and 20 to 91%, for K 29 to 100% and 20 to 57%, for Na 3 to 100% and 1 to 54%, and for Ca 0 to 32% and 0 to 30%. The species differed significantly with respect to their efficiency in nutrient resorption. Such interspecific differences in leaf nutrient economy enhance the conservative utilization of soil nutrients by the dry forest community. This reflects an adaptational strategy of the species growing on seasonally dry, nutrient‐poor soils as they tend to depend more or less on efficient internal cycling and, thus, utilize the retranslocated nutrients for the production of new foliage biomass in summer when the availability of soil moisture and nutrients is severely limited.     

重庆石灰岩地区主要木本植物叶片性状及养分再吸收特征

以重庆石灰岩地区15种常绿木本植物和14种落叶木本植物为研究对象,对两种生活型植物叶片衰老前后叶干物质含量(LDMC)、比叶面积(SLA)和叶片厚度(LT)进行了比较,并采用不同的计算方法(单位质量叶片养分含量、单位面积叶片养分含量)分析了两类植物叶片衰老前后养分含量及再吸收特征,最后对养分再吸收效率与其他叶性状因子之间的关系进行了相关分析。结果表明:常绿植物成熟叶LDMC、LT及衰老叶LT显著低于落叶植物,落叶植物成熟叶和衰老叶SLA均显著高于常绿植物(P0.05);基于单位质量叶片计算的养分含量,常绿植物成熟和衰老叶N、P量均低于落叶植物,而基于单位面积叶片计算的N、P含量则表现出相反的趋势;基于不同方法计算的N、P再吸收效率差异不明显,其中常绿植物基于单位质量叶片养分含量计算的N、P平均再吸收效率为39.42%、43.79%,落叶植物的为24.08%、33.59%;常绿和落叶植物N、P再吸收效率与LDMC、SLA、LT和成熟叶N、P含量之间没有显著相关性,但与衰老叶养分含量存在显著负相关(P0.05)。研究发现,无论是常绿植物还是落叶植物,衰老叶N、P含量均较低,表明石灰岩地区植物具有较高的养分再吸收程度。     

Canopy development of a determinate and an indeterminate cultivar of Vicia faba L. under contrasting plant distributions and densities

Field experiments were conducted in 1987 and 1988 to quantify differences in canopy formation between an indeterminate and a determinate genotype of Vicia faba L., grown at two plant densities and three spatial distributions. The number of stems per unit area produced by determinate plants was related to the growth rate before flowering. Leaf production per stem per unit of thermal time was similar in both plant types, but twice as many leaves per stem were produced by the indeterminate cultivar. The indeterminate cultivar produced fewer and smaller leaves in the warmer and drier weather of 1988 than in 1987. The determinate genotype produced similar sizes and numbers of leaves in both years, but fewer tillers developed in 1988 than in 1987. Accordingly, leaf mass per unit ground area was greater in 1987 than in 1988 in both genotypes. Except during early flowering, relationships between leaf mass and leaf area were constant, with higher specific leaf areas in the determinate than the indeterminate genotype. Shoot dry matter partitioning into leaves was identical in both years for indeterminate plants, but differed in determinate ones.
It is concluded that canopy development is regulated through individual leaf weight and leaf number per stem in non-tillering indeterminate, and by stem numbers per unit area in tillering determinate plants.     

Leaf nutrient dynamics and nutrient resorption: a comparison between larch plantations and adjacent secondary forests in Northeast China

Aims Conversion of secondary forests to pure larch plantations is a common management practice driven by the increasing demand for timber production in Northeast China, resulting in a reduction in soil nutrient availability after a certain number of years following conversion. Nutrient resorption prior to leaf senescence was related to soil fertility, an important nutrient conservation strategy for plants, being especially significant in nutrient-poor habitats. However, the seasonal dynamics of leaf nutrients and nutrient resorption in response to secondary forest conversion to larch plantations is not well understood.Methods A comparative experiment between larch plantations (Larix spp.) and adjacent secondary forests (dominant tree species including Quercus mongolica, Acer mono, Juglans mandshurica and Fraxinus rhynchophylla) was conducted. We examined the variations in leaf nutrient (macronutrients: N, P, K, Ca and Mg; micronutrients: Cu and Zn) concentrations of these tree species during the growing season from May to October in 2013. Nutrient resorption efficiency and proficiency were compared between Larix spp. and the broadleaved species in the secondary forests.Important findings Results show that the seasonal variation of nutrient concentrations in leaves generally exhibited two trends, one was a downward trend for N, P, K, Cu and Zn, and another was an upward trend for Ca and Mg. The variations in foliar nutrient concentrations were mainly controlled by the developmental stage of leaves rather than by tree species. Resorption of the observed seven elements varied among the five tree species during leaf senescence. Nutrient resorption efficiency varied 6–75% of N, P, K, Mg, Cu and Zn, while Ca was not retranslocated in the senescing leaves of all species, and Mg was not retranslocated in Larix spp. Generally, Larix spp. tended to be more efficient and proficient (higher than 6–30% and 2–271% of nutrient resorption efficiency and resorption proficiency, respectively) in resorbing nutrients than the broadleaved species in the secondary forests, indicating that larch plantations had higher leaf nutrient resorption and thus nutrient use efficiency. Compared with Larix spp., more nutrients would remain in the leaf litter of the secondary forests, indicating an advantage of secondary forests in sustaining soil fertility. In contrast, the larch plantation would reuse internal nutrients rather than lose nutrients with litter fall and thus produce a positive feedback to soil nutrient availability. In summary, our results suggest that conversion from secondary forests to pure larch plantations would alter nutrient cycling through a plant-mediated pathway.     

Evaluating a sustainability index for nutrients in a short rotation energy cropping system

In dryland environments 3–5 year rotations of tree crops and agriculture represent a major potential bioenergy feedstock and a means to restore landscape hydrologic balances and phytoremediate sites, while maintaining food production. In soils with low natural fertility, the long‐term viability of these systems will be critically affected by site nutrient status and subsequent cycling of nutrients. A nutrient assimilation index (NAI) was developed to allow comparison of species and tree component nutrient assimilation and to optimize nutrient management, by quantifying different strategies to manage site nutrients. Biomass, nutrient export and nutrient use efficiency were assessed for three short rotation tree crop species. Nutrient exports following harvest at 3 years of high density (4000 trees ha^?1) were consistently higher in Pinus radiata, with values of 85 kg ha^?1 of N, 11kg ha^?1 of P, and 62 kg ha^?1 of K, than Eucalyptus globulus and Eucalyptus occidentalis. Component NAI was generally in the order of leaf?1 for N in leaves of P. radiata to 4.7 Mg kg^?1 for P in stem‐wood of E. occidentalis, indicating higher sustainability of wood biomass compared with leaf biomass. The leaves for each species contained between 40 and 60% of the total nutrient contents while comprising around 25–30% of the total biomass. These nutrient exports via biomass removal are similar to those that follow 3 years of wheat production in the same region, indicating there is no additional drawdown of nutrient reserves during the tree cropping phase of the rotation.     

Quercus pubescens and its hemiparasite Loranthus europaeus: nutrient dynamics of leaves and twigs

Despite a long history of observations of the hemiparasitic plant, mistletoe, the mechanism of mineral movement from the host to the mistletoe is still not fully understood. In this article, we focused on the leaf development and nutrient dynamics of Loranthus europaeus and the host tree, Quercus pubescens. The nitrogen, potassium and calcium contents of leaves, current-year twigs and 1-year-old twigs were analysed. The timing of the leaf development differed between species. Leaf expansion occurred in the first 23 and 136?days, and leaf senescence took 78 and 24?days for Quercus and Loranthus, respectively. The similar nitrogen concentrations per unit leaf area may indicate that both species have the same assimilation rate. The differences in nutrient accumulation seem to support the hypothesis that nitrogen is the limiting nutrient in the transpiration stream. Larger differences in the nutrient dynamics between species were revealed in the accumulation potassium and calcium. Nutrients seemed to be transferred passively through the xylem sap between Loranthus and Quercus as we found a strong correlation between the calcium and potassium concentrations within the species and between the species. There was no correlation in the case of 1-year-old twigs, possibly due to the relatively small amount of nutrients incorporated into 1-year-old twigs and the fact that nutrient translocation occurs according the needs of the physiologically more active leaves and current-year-old twigs.     

Detrital nutrient content and leaf species differentially affect growth and nutritional regulation of detritivores

Resource nutrient content and identity are common bottom–up controls on organismal growth and nutritional regulation. One framework to study these factors, ecological stoichiometry theory, predicts that elevated resource nitrogen (N) and phosphorus (P) contents enhance organism growth by alleviating constraints on N and P acquisition. However, the regulatory mechanisms underlying this response – including whether responses depend on resource identity – remain poorly understood. In this study, we tested roles of detrital N and P contents and identity (leaf species) in constraining growth of aquatic invertebrate detritivores. We synthesized results from seven detritivore species fed wide nutrient gradients of oak and maple detritus in the laboratory. Across detritivore taxa, we used a meta‐analytic approach quantifying effects of detrital leaf species and N and P contents on growth, consumption, and N‐ and P‐specific assimilation and growth efficiencies. Detritivore growth rates increased on higher‐N and P detritus and on oak compared to maple detritus. Notably, the mechanisms of improved growth differed between the responses to detrital nutrients versus leaf species, with the former driven by greater consumption rates despite lower assimilation efficiencies on higher‐nutrient detritus, and the latter driven by improved N and P assimilation and N growth efficiencies on oak detritus. These findings suggest animal nutrient acquisition changes flexibly in response to resource changes, altering the fate of detrital N and P throughout regulation. We affirm resource identity and nutrients as important bottom–up controls, but suggest these factors act through separate pathways to affect organism growth and thereby change detrital ecosystems under anthropogenic forest compositional change and nutrient enrichment.     

Limiting effects of low leaf-water content on the nitrogen utilization,energy budget,and larval growth ofHyalophora cecropia (Lepidoptera: Saturniidae)

Summary Hyalophora cecropia larvae were reared on leaves of wild cherry,Prunus serotina, which contained variable amounts of leaf water but otherwise did not differ in fiber, total nitrogen, and caloric content. Larvae which were fed leaves low in leaf water grew more slowly and were less efficient at utilizing plant biomass, energy, and nitrogen than those larvae fed leaves which were fully supplemented with water.Experiments were performed using excised leaves under different regimes of relative humidity and leaf water supplementation in climatic control chambers maintained at identical temperatures and photoperiod. Foodplant biomass utilization efficiencies were severely reduced by decreasing amounts of leaf water. Growth rates were halved and the efficiency of conversion of assimilated dry matter into larval biomass was reduced from 82% in the treatment with fully supplemented leaves to 34% in the driest treatment. The nitrogen utilization efficiency (N.U.E.) was reduced from 75–80% to 48%, and the relative accumulation rate of nitrogen (N.A.R.) was suppressed nearly 2-fold for larvae on low-water leaves. Relative maintenance costs (calories expended in respiration/mg tissue/day) of larvae were nearly five times higher on dry leaves than on fully supplemented leaves. Larvae on leaves which were low in water content were themselves more desiccated, and metabolized greater portions of assimilated energy, perhaps in an attempt to supplement body water with metabolic water derived from respiration.The larval rates of consumption of biomass, energy, and nitrogen were the same for all treatments, indicating that leaf water affected larval growth primarily by restricting the efficiency of utilizing these nutrients. Where water was limiting (as in tree leaves), an increased consumption rate did not appear to be a successful means of increasing growth rates. There were daily and seasonal differences in leaf water content between different trees of the same species. Although absolute differences in leaf water exist between different trees and between young and old (fully expanded) leaves of a single tree, these differences are proportional and parallel each other through daily and seasonal cycles.In spite of evolutionary adaptations of herbivores to acquire adequate water and avoid desiccation, the leaf water content naturally encountered by cecropia larvae on cherry leaves may limit their growth, especially if the R.H. is low.     

Communication of leaf suitability by gregarious eastern tent caterpillars (Malacosoma americanum)

ABSTRACT. 1. Previous work has shown that leaf age affects recruitment trail marking by eastern tent caterpillars ( Malacosoma americanum Fabr.). Young leaves of host plants elicit trail marking to a greater degree than mature leaves.
2. Experiments were conducted to establish the relationship between the differential behavioural responses of larvae to young and mature leaves and the suitability of foliage for larval growth and survival. Foliage of black cherry ( Prunus serotina Ehrh.), a typical rosaceous host plant, was used for this comparison.
3. Larvae preferred young leaves to mature leaves in choice tests, and marked more to young leaves than to mature leaves in no-choice tests.
4. Mature leaves supported adequate growth through two larval instars of rearing, but thereafter were unsuitable for growth. Larvae fed mature leaves had lower pupal weight, poorer survival, and grew less efficiently than larvae fed young leaves.
5. The results support the hypothesis that the trail communication system of eastern tent caterpillars is an adaptation to efficiently locate leaves which are favourable for larval growth and survival.     

Chloroplastic ATP synthase optimizes the trade-off between photosynthetic CO2 assimilation and photoprotection during leaf maturation

In the present study, we studied the role of chloroplastic ATP synthase in photosynthetic regulation during leaf maturation. We measured gas exchange, chlorophyll fluorescence, P700 redox state, and the electrochromic shift signal in mature and immature leaves. Under high light, the immature leaves displayed high levels of non-photochemical quenching (NPQ) and P700 oxidation ratio, and higher values for proton motive force (pmf) and proton gradient (ΔpH) across the thylakoid membranes but lower values for the activity of chloroplastic ATP synthase (g_H⁺) than the mature leaves. Furthermore, g_H⁺ was significantly and positively correlated with CO₂ assimilation rate and linear electron flow (LEF), but negatively correlated with pmf and ΔpH. ΔpH was significantly correlated with LEF and the P700 oxidation ratio. These results indicated that g_H⁺ was regulated to match photosynthetic capacity during leaf maturation, and the formation of pmf and ΔpH was predominantly regulated by the alterations in g_H⁺. In the immature leaves, the high steady-state ΔpH increased lumen acidification, which, in turn, stimulated photoprotection for the photosynthetic apparatus via NPQ induction and photosynthetic control. Our results highlighted the importance of chloroplastic ATP synthase in optimizing the trade-off between CO₂ assimilation and photoprotection during leaf maturation.     

Does the Gradualness of Leaf Shedding Govern Nutrient Resorption from Senescing Leaves in Mediterranean Woody Plants?

To reveal the environmental and substrate quality effects on decomposition process and enzyme activities, litterbag experiments containing Nuphar and Carex leaves, Nuphar rhizome, and Ranunculus shoot, were carried in five-subalpine marshes in Lake Tahoe basin, USA. Alkaline phosphatase, β-glucosidase, and β-xylosidase activities were determined by a fluorogenic method using methyumbelliferyl substrates. Carex leaves, Nuphar rhizome and leaves, and Ranunculus shoots lost, respectively, 33, 67, 82 and 93% of original dry weight over 268 days. Decay rates were different among substrates but not among marshes. Nitrogen and carbon contents increased during the first 58 days and subsequently remained stable. Phosphorus content was stable during the experimental period except for a decrease in the first 16 days in Nuphar shoots. Enzyme activities in decomposing Carex and Nuphar leaves in four marshes were not significantly affected by environmental conditions. β-glucosidase and β-xylosidase activities in decomposing Carex leaves increased with time, but in other plant tissue these enzyme activities remained stable during experimental period. Enzyme activities were significantly different among decomposing substrates. Alkaline phosphatase activity was highest in Nuphar leaves (ca. 1286 μ-mole h⁻¹ g DW −1) but lower and similar in other plant tissues (ca. 100 and 10 μ-mole h −1 g DW −1, respectively). This study showed differences in decay rates and enzyme activities rely on substrate and not the environment conditions of the study area. Decomposition rates in the early stage of decomposition were related to cumulative enzyme activities.     

Latitudinal Patterns and Climatic Drivers of Leaf Litter Multiple Nutrients in Chinese Broad-Leaved Tree Species: Does Leaf Habit Matter?

Leaf litter nutrients play a key role in nutrient cycling in forest ecosystems, yet our current knowledge of the ways in which climate controls leaf litter nutrients remains uncertain, especially for broad-leaved tree species in China. We performed a meta-analysis of geographic patterns of leaf litter nutrients of Chinese broad-leaved tree species in relation to climatic variables and leaf habit (as a discrete classification of tree species). We found that mean leaf litter carbon (C), nitrogen (N), phosphorus (P), potassium (K), calcium (Ca) and magnesium (Mg) were 458.36, 10.11, 0.72, 6.37, 14.22 and 2.59 mg^?1 g, respectively. Leaf litter nutrients did not diverge between leaf habits where they coexisted. These leaf litter nutrients displayed significant latitudinal trends, partly driven by climatic factors and a shift in leaf habit. Mean annual precipitation explained the largest amount of total variation in leaf litter C, N, P and K, and mean annual temperature was the most important predictor for leaf litter Mg, whereas leaf habit was the largest contributor to total variation in leaf litter Ca. We further found that the relationships between climate and leaf litter nutrients were distinguishable for evergreen and deciduous broad-leaved tree species. Collectively, our study differed from previous studies that evaluated leaf litter nutrients and only focused on N and P, and substantiated that leaf litter nutrients in forest ecosystems were affected by climate and leaf habit, but the strengths of the influences of these factors were strongly contingent on leaf litter nutrient identity. Therefore, alteration of climate would directly and indirectly (via a shift in species composition) affect latitudinal patterns of leaf litter nutrients and thus the associated nutrient flux and ecosystem functioning. Our study also underlined the need to include multiple nutrients to explore the influence of climate on leaf litter nutrient stoichiometry.     

Positive indirect effect of tadpoles on a detritivore through nutrient regeneration

In aquatic food webs consumers can affect other members of the web by releasing nutrients as a result of their feeding activity. There is increasing evidence of these positive effects on primary producers, but such nutrient regeneration can also affect detritivores, by favoring the activities of detritus-associated microbes. We examined the effects of nutrient regeneration by tadpoles on leaf-eating detritivores under laboratory conditions. We fed four species of tadpoles three different food items (leaf litter, algae, and sludgeworms). We then conditioned terrestrial dead leaves with water from reared tadpoles (treatments) or food items alone (controls), and compared the C:N ratios of the conditioned leaves and the growth of the isopod Asellus hilgendorfii fed on the conditioned leaves. Tadpole feeding activity reduced the C:N ratio of conditioned leaves, and the effect was greatest when tadpoles were fed algae. Isopod growth rates were often higher when they were fed the litter conditioned with water from reared tadpoles. Thus, nutrient regeneration by tadpoles had a positive indirect effect on detritivores by enhancing leaf quality. Tadpoles often occur in nutrient-limited habitats where leaf litter is the major energy source, and their facilitative effects on leaf-eating detritivores may be of great significance in food webs by enhancing litter decomposition.