Temperature sensitivity of nitrogen productivity for Scots pine and Norway spruce

Environmental conditions control physiological processes in plants and thus their growth. The predicted global warming is expected to accelerate tree growth. However, the growth response is a complex function of several processes with both direct and indirect effects. To analyse this problem we have used needle nitrogen productivity, which is an aggregate parameter for production of new foliage. Data on needle dry matter, production, and nitrogen content in needles of Scots pine ( Pinus sylvestris) and Norway spruce ( Picea abies) from a wide range of climatic conditions were collected and needle nitrogen productivities, defined as dry matter production of needles per unit of nitrogen in the needle biomass, were calculated. Our results show that the nitrogen productivity for spruce is insensitive to temperature. However, for pine, temperature affects both the magnitude of nitrogen productivity at low needle biomass and the response to self-shading but the temperature response is small at the high end of needle biomass. For practical applications it may be sufficient to use a species-specific nitrogen productivity that is independent of temperature. Because temperature affects tree growth indirectly as well as through soil processes, the effects of temperature change on tree growth and ecosystem carbon storage should mainly be derived from effects on nitrogen availability through changes in nitrogen mineralization. In addition, this paper summarises data on dry matter, production and nitrogen content of needles of conifers along a temperature gradient.     

鼎湖山某些植物群落根系生物量及其氮素动态

 本文用挖土柱及埋土柱法测定了鼎湖山生物圈保护区季风常绿阔叶林,黄果厚壳桂、鼎湖钓樟群落的根系生物量和年生长量。用自动离子分析仪分析了根的含氮量并研究了根系的氮素动态。结果表明,根系生物量(t／ha)为35.43,其中活根26.63,死根8.8;根的生长量(kg／ha·a)是3.74。根库贮氮量(kg／ha)为279.23,其中活根库为202.25,死根库为76.98;每年新生根系积累的氮素为57.55kg／ha。每年新生根系生产量及其氮素积累量占整个根库生物量及其氮素贮量的百分率分别为10.6及20.6,根库年生长量及其氮素积累的比率均较高,群落正处在生长发育盛期。     

The influence of nitrogen availability on carbon and nitrogen storage in the biennial Cirsium vulgare (Savi) Ten. II. The cost of nitrogen storage

The cost of nitrogen storage to current growth was examined in relation to N availability in the biennial Cirsium vulgare. Plants were grown outdoors, in sand culture, with continuous diel drip irrigation of fertilization medium containing one of five different N concentrations. Plants grown at the highest N concentration stored twice as much N in their tap roots as did plants grown at the lowest N concentration. In high-N-grown plants, the storage of N reserves occurred during the period of maximum growth, at the same time as tap-root production. At the time of maximum biomass, stored N was also at a maximum. During the period following maximum biomass, no additional storage of N occurred. This pattern was observed despite frequent late-season leaf senescence which resulted in a large pool of potentially mobile N which could have been stored at no cost to growth. In low-N-grown plants, the production of tap-root storage tissue and the filling of that tissue with stored N were staggered. Tap-root production and growth occurred during the period of maximum growth, as in the high-N-grown plants. However, filling of the storage tissue with N occurred late in the growing season, when the pool of mobile N from senescent leaves was large. The utilization of this late-season N source occurred with little or no cost to growth, and this N is labelled, according to previous definitions, as ‘accumulated’. The costs of storing N in plants of the different N treatments were calculated using two models based on different growth constraints. In one model, the cost of N storage was represented as lost growth due to allocation of N to storage, rather than to the photosynthetic shoot (i.e. growth was assumed to be limited by carbon acquisition). In the second model, the storage cost was calculated as lost growth due to allocation of N to storage, rather than to the nitrogen-acquiring fine-root system (i.e. growth was assumed to be limited by nitrogen acquisition). In both models, the total cost of N storage was predicted to decrease as N availability decreased due to smaller storage pool sizes in plants of the low-N treatments. The cost of filling the tap root with stored N as a percentage of the total storage cost was also reduced as N availability decreased due to the occurrence of late-season accumulation. By relying, at least in part, on late-season accumulation, plants grown at the lowest three levels of N availability reduced total storage costs by 15 to 22%. The results demonstrate that plants are capable of adjusting their storage patterns in response to low nitrogen availability such that the costs of storage are reduced.     

Effects of experimental warming on growth,biomass allocation,and needle chemistry of <Emphasis Type="Italic">Abies faxoniana</Emphasis> in even-aged monospecific stands

The response and adaption mechanisms of seedlings under long-term warming have remained largely unknown. In this study, we investigated the effects of warming for 6 years on growth, and needle carbon, nitrogen, chlorophyll, and carbohydrate levels in a coniferous tree species, Abies faxoniana. Seedlings were grown in even-aged monospecific stands under ambient and warming (ambient +2.2°C) temperature in climate control chambers. Warming caused statistically significant increases in the specific leaf area, leaf area ratio, root biomass, leaf biomass, branch biomass, stem biomass, and total mass of the seedlings, and reduced the root/shoot ratio. Warming also increased total chlorophyll concentrations, specific chlorophyll pigments, and Chlorophyll a/b ratios in both studied needle age classes. In addition, C/N ratios of current-year and 1-year-old needles increased by warming. In contrast, warming decreased the levels of N, sugar, cellulose, and starch in needles, while warming had no effect on the height, stem diameter, needle mass ratio, root mass ratio, and root/needle ratio. We conclude that warming increases branch growth and changes needle chemistry, which enhances the light capture potential of seedlings.     

光照和氮素对喜旱莲子草形态特征和生物量分配的影响

研究了两个光照梯度和3个土壤氮素水平交互作用对喜旱莲子草(Alternan thera philoxeroides(Mart.)Griseb.)形态特征和生物量分配的影响。结果表明,全光照促进喜旱莲子草总生物量的积累,但在遮荫条件下,喜旱莲子草可以通过增加株高、光合叶面积和改变生物量分配来适应弱光生境。土壤中氮素含量对喜旱莲子草生长有明显影响,总生物量、株高、叶面积、茎生物量比和叶生物量比等随土壤氮素水平增加而增加。光照和氮素的交互作用对总生物量、根生物量比、茎生物量比和叶生物量比也有显著影响。随着氮素水平的增高,遮荫和高光照处理下喜旱莲子草的叶面积、总生物量和叶生物量比之间的差异减小,而株高和根生物量比之间的差异增大。此外,光照强度对茎生物量比的影响具有明显的氮素浓度依赖性,低氮条件下,茎生物量比在高光照处理下显著高于遮荫处理,而在中氮条件下,遮荫处理却显著高于高光照处理,且在高氮处理下其差异进一步加大。这些结果表明喜旱莲子草在高氮素环境下能够通过形态可塑性和生物量分配模式的改变来适应弱光环境所带来的不利影响。研究结果不但可为研究喜旱莲子草对异质生境的入侵机制提供资料,也可为进一步研究喜旱莲子草的入侵和扩散与农业等生态系统中土壤氮素残留的关系提供参考。       

Nitrogen availability affects saprotrophic basidiomycetes decomposing pine needles in a long term laboratory study

Fungi, especially basidiomycetous litter decomposers, are pivotal to the turnover of soil organic matter in forest soils. Many litter decomposing fungi have a well-developed capacity to translocate resources in their mycelia, a feature that may significantly affect carbon (C) and nitrogen (N) dynamics in decomposing litter. In an eight-month long laboratory study we investigated how the external availability of N affected the decomposition of Scots pine needles, fungal biomass production, N retention and N-mineralization by two litter decomposing fungi – Marasmius androsaceus and Mycena epipterygia. Glycine additions had a general, positive effect on fungal biomass production and increased accumulated needle mass loss after 8 months, suggesting that low N availability may limit fungal growth and activity in decomposing pine litter. Changes in the needle N pool reflected the dynamics of the fungal mycelium. During late decomposition stages, redistribution of mycelium and N out from the decomposed needles was observed for M. epipterygia, suggesting autophagous self degradation.     

The species richness–biomass relationship in herbaceous plant communities: what difference does the incorporation of root biomass data make?

So far, in all studies on the much-discussed hump-backed relationship between plant community productivity and species richness, productivity has been assessed through plant shoot biomass, i.e. it has been ignored that frequently most of the biomass is produced below ground. We revisited the 27 grassland and forest field-layer communities, studied earlier by Zobel and Liira, to sample root biomass, plant total biomass and root/shoot allocation, and learn how the incorporation of below-ground biomass data would affect the shape of the hump-backed relationship. In order to avoid scaling artefacts we estimated richness as the average count of species per 500 plant ramets (absolute richness). We also included relative richness measures. Relative richness was defined as richness per 500 ramets/size of the actual species pool (the set of species present in the community), relative pool size was defined as size of the actual species pool/size of the regional species pool (the set of species available in the region and capable of growing in the given community).
The biomass-absolute richness relationship was humped, irrespective of the biomass measure used, the hump being most obvious when plant total biomass was used as the independent variable. Evidently, the unimodal richness–productivity curve is not a sampling artefact, as suspected by Oksanen. However, relative richness was not related to community biomass (above-ground, below-ground or total). The hump-backed curve is shaped by the sizes of actual species pools in communities, implying that processes which are responsible for small-scale diversity pattern mainly operate on the community level.
Neither absolute nor relative richness were significantly related to root/shoot allocation. The presumably stronger (asymmetric) shoot competition at greater allocation to shoots appears not to suppress small-scale richness. However, there is a significant relationship between relative pool size and root/shoot allocation. Relatively more species from regional species pools are able to enter and persist in communities with more biomass allocated into roots.     

Growth and Nutrient Retranslocation in Needles of Radiata Pine in Relation to Nitrogen Supply

The effects of N application on tree growth and the retranslocationof N, P, and K from young needles to new growth were examinedin young radiata pine (Pinus radiata D. Don) trees. Nitrogen fertilization increased the number and size of needles,rates of shoot production, stem volume growth and tree biomass.Foliar N and P contents (µg per needle) fluctuated ina cyclic fashion with prominent phases of accumulation, retranslocationand replenishment. The patterns of these fluctuations in controland N-fertilized trees were similar, although the fluxes ofN, P and K in and out of needles were increased by N fertilization.Greater translocation (g per tree) of N and K from needles ofN fertilized trees occurred because fertilization increasedthe needle weight and the proportion of N and K retranslocatedfrom individual needles. Nitrogen fertilization increased theretranslocation of P largely as a result of higher needle mass.Trees supplied with more than adequate amounts of P in the soilretranslocated up to 58 per cent of the initial pool of P fromyoung needles. The periods of high retranslocation coincidedwith periods of high concentrations of soil mineral N and withshoot production. Conversely, the periods of rapid replenishmentof N and P into the needles coincided with the time of slowshoot growth and low concentration of soil mineral N. The growthrate of trees, rather than the availability of nutrients inthe soil was the main factor controlling retranslocation. For radiata pine, retranslocation from needles is not a mechanismspecific for coping with low soil fertility. It seems to bea mechanism which enhances the nutrient supply to apical growingpoints, especially during periods of flushing. Pinus radiata, nitrogen supply, shoot growth, nutrient fluctuations and retranslocation, nutrient use and adaptation     

General allometric equations and biomass allocation of <Emphasis Type="Italic">Pinus massoniana</Emphasis> trees on a regional scale in southern China

Applying allometric equations in combination with forest inventory data is an effective approach to use when qualifying forest biomass and carbon storage on a regional scale. The objectives of this study were to (1) develop general allometric tree component biomass equations and (2) investigate tree biomass allocation patterns for Pinus massoniana, a principal tree species native to southern China, by applying 197 samples across 20 site locations. The additive allometric equations utilized to compute stem, branch, needle, root, aboveground, and total tree biomass were developed by nonlinear seemingly unrelated regression. Results show that the relative proportion of stem biomass to tree biomass increased while the contribution of canopy biomass to tree biomass decreased as trees continued to grow through time. Total root biomass was a large biomass pool in itself, and its relative proportion to tree biomass exhibited a slight increase with tree growth. Although equations employing stem diameter at breast height (dbh) alone as a predictor could accurately predict stem, aboveground, root, and total tree biomass, they were poorly fitted to predict the canopy biomass component. The inclusion of the tree height (H) variable either slightly improved or did not in any way increase model fitness. Validation results demonstrate that these equations are suitable to estimate stem, aboveground, and total tree biomass across a broad range of P. massoniana stands on a regional scale.     

植物代谢速率与个体生物量关系研究进展

植物的各项生理生态功能（例如,呼吸、生长和繁殖）都与个体生物量成异速生长关系。West, Brown及Enquist基于分形网络结构理论所提出的WBE模型认为：植物的代谢（呼吸）速率正比于个体生物量的3/4次幂。然而,恒定的“3/4异速生长指数”与实测数据、植物生理生态学等研究之间存在矛盾,引发激烈的争论。论文分析了不同回归方法对代谢指数的影响,重点对植物代谢速率与个体生物量异速生长关系研究进展进行了综述,分析并得出了植物代谢指数在小个体时接近1.0,并随着生物量的增加而系统减小,且其密切依赖于氮含量的调控的结论。据此,提出了进一步深入研究植物代谢速率个体生物量关系需要解决的一些科学问题。     

Nitrogen distribution in young Norway spruce (Picea abies) trees as affected by pedospheric nitrogen supply

In numerous locations in Europe spruce trees are exposed to high loads of nitrogen. The present study was performed to characterize the distribution of nitrogen compounds under these conditions. For this purpose Norway spruce ( Picea abies [L.] Karst.) trees were cultivated under close-to-natural conditions of a forest understory in soil from an apparently nitrogen-limited field site in the Black Forest either with, or without supplementation of nitrogen as ammonium nitrate. After 11 and 20 months, growth, total nitrogen contents of the biomass, and total soluble non-proteinogenic nitrogen compounds (TSNN, i.e. nitrate, ammonium, soluble proteinogenic and non-proteinogenic amino compounds) in needles, xylem sap and phloem exudate were analysed. After 20 months of growth, N-fertilization had slightly enhanced the biomass of current-, but not of 1-year-old shoots. At both harvests, total N-content of 1-year-old needles was increased by N-fertilization, whereas current-year needles were not significantly affected. By contrast, TSNN was elevated by N-fertilization in both current-year and 1-year-old needles. The increase in TSNN was mainly attributed to an accumulation of arginine. Xylem sap analysis showed that the increase in TSNN of the needles was a consequence of enhanced nitrogen assimilation of the roots rather than the shoot. Since also TSNN in phloem exudates was enhanced, it appears that N-fertilization elevates the cycling pool of amino compounds in young Norway spruce trees. However, this pool seems to be subject to metabolic interconversion, since mainly glutamine and aspartate are transported in the xylem from the roots to the shoot, but arginine accumulated in the needles and the phloem.     

Effects of phosphorus supply on growth and nitrogen fractions in xylem sap and foliage of Eucalyptus regnans (F.Muell.), E. nitens (Maiden) and E. globulus (Labill.) seedlings: implications for herbivory

Rates of growth of seedlings of E. globulus, E. regnans and E. nitens were related to phosphorus supply in two soils but concentrations of total nitrogen and total phosphorus in most plant tissues did not vary significantly among soil or phosphorus treatments. Differences in concentrations of nitrogen and phosphorus and in the composition of the pool of free amino-acids among leaves at different stages of development were far greater than differences between treatments. The most significant of these differences were several-fold greater concentrations of arginine in the oldest leaves and these are most likely due to protein degradation and/or in situ synthesis since arginine is not generally phloem mobile. The concentration of reduced nitrogen in xylem sap was inversely related to growth and glutamine was by far the dominant nitrogenous solute. We suggest that specific nitrogenous solutes may be useful indices of the nitrogen status of eucalypt tissues for insect herbivores.     

Growth and reproduction of Arabidopsis thaliana in relation to storage of starch and nitrate in the wild-type and in starch-deficient and nitrate-uptake-deficient mutants

We have investigated the interactions between resource assimilation and storage in rosette leaves, and their impact on the growth and reproduction of the annual species Arabidopsis thaliana. The resource balance was experimentally perturbed by changing (i) the external nutrition, by varying the nitrogen supply; (ii) the assimilation and reallocation of resources from rosette leaves to reproductive organs, by cutting or covering rosette leaves at the time of early flower bud formation, and (iii) the internal carbon and nitrogen balance of the plants, by using isogenic mutants either lacking starch formation (PGM mutant) or with reduced nitrate uptake (NU mutant). When plants were grown on high nitrogen, they had higher concentrations of carbohydrates and nitrate in their leaves during the rosette phase than during flowering. However, these storage pools did not significantly contribute to the bulk flow of resources to seeds. The pool size of stored resources in rosette leaves at the onset of seed filling was very low compared to the total amount of carbon and nitrogen needed for seed formation. Instead, the rosette leaves had an important function in the continued assimilation of resources during seed ripening, as shown by the low seed yield of plants whose leaves were covered or cut off. When a key resource became limiting, such as nitrogen in the NU mutants and in plants grown on a low nitrogen supply, stored resources in the rosette leaves (e.g. nitrogen) were remobilized, and made a larger contribution to seed biomass. A change in nutrition resulted in a complete reversal of the plant response: plants shifted from high to low nutrition exhibited a seed yield similar to that of plants grown continuously on a low nitrogen supply, and vice versa. This demonstrates that resource assimilation during the reproductive phase determines seed production. The PGM mutant had a reduced growth rate and a smaller biomass during the rosette phase as a result of changes in respiration caused by a high turnover of soluble sugars ( Caspar et al. 1986 ; W. Schulze et al. 1991 ). During flowering, however, the vegetative growth rate in the PGM mutant increased, and exceeded that of the wild-type. By the end of the flowering stage, the biomass of the PGM mutant did not differ from that of the wild-type. However, in contrast to the wild-type, the PGM mutant maintained a high vegetative growth rate during seed formation, but had a low rate of seed production. These differences in allocation in the PGM mutant result in a significantly lower seed yield in the starchless mutants. This indicates that starch formation is not only an important factor during growth in the rosette phase, but is also important for whole plant allocation during seed formation. The NU mutant resembled the wild-type grown on a low nitrogen supply, except that it unexpectedly showed symptoms of carbohydrate shortage as well as nitrogen deficiency. In all genotypes and treatments, there was a striking correlation between the concentrations of nitrate and organic nitrogen and shoot growth on the one hand, and sucrose concentration and root growth on the other. In addition, nitrate reductase activity (NRA) was correlated with the total carbohydrate concentration: low carbohydrate levels in starchless mutants led to low NRA even at high nitrate supply. Thus the concentrations of stored carbohydrates and nitrate are directly or indirectly involved in regulating allocation.     

Nutrient limitation and plant species composition in temperate salt marshes

Addition of inorganic nitrogen, phosphorus and potassium in a factorial design in two ungrazed Wadden-Sea salt marshes at low and high elevations showed that nitrogen was the limiting nutrient. No effects of nutrient addition were detected in the 1st year, probably due to a considerable rainfall deficit during the growing season. In the 2nd year, which was more humid, only nitrogen addition caused significant effects in both the low salt marsh dominated by Puccinellia maritima and the high marsh dominated by Festuca rubra. No two-way or three-way interactions with phosphorus or potassium were found. In the low marsh, nitrogen addition had a negative effect on the biomass of Puccinellia, but a positive effect on the biomass of Suaeda maritima and on the total above-ground biomass. Puccinellia was replaced by Suaeda after nitrogen addition, due to shading. In the high salt marsh, no significant effects of fertilizer application on total above-ground biomass were found, due to the weak response of the dominant species Festuca rubra, which accounted for 95% of total biomass. The biomass of Spergularia maritima increased, however, as a response to nitrogen addition.The shoot length of Festuca was positively affected by nitrogen fertilization. It is suggested that stands of Festuca reached maximal biomass at the study site without fertilization and that its growth was probably limited by self-shading. Received: 22 September 1996 / Accepted: 5 April 1997     

Microbial biomass C,N and P in two arctic soils and responses to addition of NPK fertilizer and sugar: implications for plant nutrient uptake

The soil microbial carbon (C), nitrogen (N) and phosphorus (P) pools were quantified in the organic horizon of soils from an arctic/alpine low-altitude heath and a high-altitude fellfield by the fumigation-extraction method before and after factorial addition of sugar, NPK fertilizer and benomyl, a fungicide. In unamended soil, microbial C, N and P made up 3.3–3.6%, 6.1–7.3% and 34.7% of the total soil C, N and P content, respectively. The inorganic extractable N pool was below 0.1% and the inorganic extractable P content slightly less than 1% of the total soil pool sizes. Benomyl addition in spring and summer did not affect microbial C or nutrient content analysed in the autumn. Sugar amendments increased microbial C by 15 and 37% in the two soils, respectively, but did not affect the microbial nutrient content, whereas inorganic N and P either declined significantly or tended to decline. The increased microbial C indicates that the microbial biomass also increased but without a proportional enhancement of N and P uptake. NPK addition did not affect the amount of microbial C but almost doubled the microbial N pool and more than doubled the P pool. A separate study has shown that CO₂ evolution increased by more than 50% after sugar amendment and by about 30% after NPK and NK additions to one of the soils. Hence, the microbial biomass did not increase in response to NPK addition, but the microbes immobilized large amounts of the added nutrients and, judging by the increased CO₂ evolution, their activity increased. We conclude: (1) that microbial biomass production in these soils is stimulated by labile carbon and that the microbial activity is stimulated by both labile C and by nutrients (N); (2) that the microbial biomass is a strong sink for nutrients and that the microbial community probably can withdraw substantial amounts of nutrients from the inorganic, plant-available pool, at least periodically; (3) that temporary declines in microbial populations are likely to release a flush of inorganic nutrients to the soil, particularly P of which the microbial biomass contained more than one third of the total soil pool; and (4) that the mobilization-immobilization cycles of nutrients coupled to the population dynamics of soil organisms can be a significant regulating factor for the nutrient supply to the primary producers, which are usually strongly nutrient-limited in arctic ecosystems.     

Competition between co-dominant plants of the Serengeti plains depends on competitor identity,water, and urine

Kyllinga nervosa (Steud.) and Sporobolus kentrophyllus (K. Schum.) are co-dominant plants of the Serengeti short-grass plains, Tanzania. The plains are characterized by seasonal and sporadic rainfall and currently support in excess of 1.5 million migratory ungulates. The interactive effect of simulated bovine urine and water availability were tested on the competitive interactions of these species in the laboratory. Sporobolus kentrophyllus was a superior competitor to K. nervosa over the tested treatment levels with respect to growth and reproductive effort. Sporobolus kentrophyllus exhibited rapid growth in response to urine addition, leading to a significant species × urine interaction while reduced growth by K. nervosa in response to low water availability explained the significant species × water interaction and is likely explained by K. nervosa's shallow root system. Kyllinga nervosa, however, appears to be more tolerant of low nitrogen conditions based on its similar growth with and without the urine treatment. The effect of intraspecific competition on total biomass was similar for S. kentrophyllus and K. nervosa. Competition resulted in increased size differences (asymmetry) for K. nervosa and for the interspecific competition treatments compared to the size differences observed for plants grown individually (in absence of competition).Total reproductive biomass was reduced most by competition with S. kentrophyllus, irrespective of target species. The water treatment did not influence reproduction while the urine treatment significantly increased reproductive biomass and interacted with target species, competitor species, and yielded a three-way urine × target × competitor species interaction.Results suggest that codominance of these two species in the Serengeti is regulated by water availability, nitrogen input from grazers, and local neighbor identity.     

秦岭华山松营养诊断与林地施肥

以陕西秦岭南坡华山松天然林为对象,采集华山松针叶、凋落物、土壤样品,测定全氮(TN)和全磷(TP)含量,开展华山松林木营养诊断,分析华山松林养分限制格局,研究促进华山松正常生长的施肥种类和施肥量,为林地精准施肥和科学培育华山松大径材提供支持。结果表明:华山松大径材率(胸径DBH>26 cm)为29.0%,具有成长为大径材潜力的华山松个体(18 cm-1,N∶P为10.24(<14的林木生长不受限阈值);N、P回收效率分别为33.8%和48.0%。在0~10 cm土层,土壤与华山松针叶的N、P含量呈显著负相关,凋落物与土壤的N、P含量呈显著正相关,土壤N储量与华山松针叶N∶P呈显著二次函数关系。华山松正常生长受土壤N限制,在林地补充0.42 t N·hm^-2可解除华山松生长限制。为快速培育大径材,宜向林地内具有成长为大径材潜力的个体施0.16 t N·hm^-2,按单位面积内林木个体生物量比例进行单株氮素分配。     

Nitrogen resorption from senescing leaves of three salt marsh plant species

Seasonalvariation in leaf nitrogen of mature green and senescent leaves and nitrogenresorption efficiency in three plants (Spartina maritima, Halimioneportulacoides and Arthrocnemum perenne) of aTagus estuary salt marsh are reported. Total nitrogen concentrations in greenand senescent leaves were higher during winter (December and March). Soilinorganic nitrogen availability showed an opposite pattern with higherconcentrations during summer (June and September) when total leaf biomass washigher. Nitrogen resorption efficiency ranged between 31 and 76% andH. portulacoides was the plant that better minimizednitrogen loss by this process. Nitrogen resorption occurred mainly from thesoluble protein pool, although other fractions must have been broken down duringthe resorption process. No significant seasonal variation in nitrogen resorptionefficiency and no relation to leaf total nitrogen or soil nitrogen availabilitywere found. This suggests that the efficiency of the resorption process is notdetermined by the plant nitrogen status nor by the availability of the nutrientin the soil. Nevertheless, resorption from senescing leaves may play animportant role in the nitrogen dynamics of salt marsh plants and reduce thenitrogen requirements for plant growth.     

Biomass loss and change in species dominance shift stream community excretion stoichiometry during severe drought

1. Animals contribute significantly to nutrient cycling through excretion, but most studies consider their effects under relatively benign abiotic conditions. Disturbances such as drought may alter animals’ nutrient contributions through shifts in species composition and biomass. Headwater streams are particularly vulnerable to extreme climate events and thus might show rapid changes in stream biota and their ecosystem effects.
2. We tested how biomass and subsequent ecosystem effects (nutrient cycling) of an intermittent prairie stream community changed during a drought. We quantified the biomass and contributions to nutrient cycling for assemblages comprising fishes, crayfish, and tadpoles in 12 isolated pools over 3 months encompassing the harshest drought on record for Kings Creek, KS, U.S.A. We predicted that macroconsumer biomass would decline with pool surface area and that differences in macroconsumer biomass and taxonomic composition would lead to different contributions of pool assemblages to nutrient cycling.
3. The biomass of pool assemblages declined with decreasing pool size, a pattern apparently driven by mortality, emigration, or metamorphosis. We also observed a change in assemblage structure of drying pools during drought relative to pool size, shifting dominance toward species with more drought-resistant traits. Accordingly, assemblage nitrogen (N) excretion rates declined as pool biomass was reduced, leading to a 58% reduction in N available to epilithic biofilms. Phosphorus (P) excretion rates declined from June to July, but increased in August, as species with high P excretion rates maintained similar proportional biomass and biomass of a non-native fish increased. Molar N:P of pool assemblage excretion declined significantly throughout the drought and coincided with loss of southern redbelly dace (Chrosomus erythrogaster: Cyprinidae).
4. Animal-mediated nutrient cycling was altered by the loss of biomass and stoichiometric traits of taxa that differed in their occurrences and ability to tolerate abiotic conditions during drought. Elevated availability of dissolved N in isolated pools may increase N uptake rates by biofilms during drought conditions, indicating the importance of N excreted by aggregated macroconsumers, especially those with unique stoichiometric traits. While the significance of shifts in the composition of freshwater communities to ecosystems is not entirely known, additional losses in ecosystem function and changes in community structure may follow episodes of severe drought.