Strategies and agronomic interventions to improve the phosphorus-use efficiency of farming systems

Phosphorus (P)-deficiency is a significant challenge for agricultural productivity on many highly P-sorbing weathered and tropical soils throughout the world. On these soils it can be necessary to apply up to five-fold more P as fertiliser than is exported in products. Given the finite nature of global P resources, it is important that such inefficiencies be addressed. For low P-sorbing soils, P-efficient farming systems will also assist attempts to reduce pollution associated with P losses to the environment. P-balance inefficiency of farms is associated with loss of P in erosion, runoff or leaching, uneven dispersal of animal excreta, and accumulation of P as sparingly-available phosphate and organic P in the soil. In many cases it is possible to minimise P losses in runoff or erosion. Uneven dispersal of P in excreta typically amounts to ~5% of P-fertiliser inputs. However, the rate of P accumulation in moderate to highly P-sorbing soils is a major contributor to inefficient P-fertiliser use. We discuss the causal edaphic, plant and microbial factors in the context of soil P management, P cycling and productivity goals of farms. Management interventions that can alter P-use efficiency are explored, including better targeted P-fertiliser use, organic amendments, removing other constraints to yield, zone management, use of plants with low critical-P requirements, and modified farming systems. Higher productivity in low-P soils, or lower P inputs in fertilised agricultural systems can be achieved by various interventions, but it is also critically important to understand the agroecology of plant P nutrition within farming systems for improvements in P-use efficiency to be realised.     

Impact of soil tillage on the robustness of the genetic component of variation in phosphorus (P) use efficiency in barley (Hordeum vulgare L.)

To enhance the sustainability of agriculture it is imperative that the use of P-fertilisers by temperate cereal crops be improved. This can be achieved both by agronomic and genetic approaches. While many studies have demonstrated genotypic variation in P-use efficiency in a number of cereal species the robustness of this genetic variation in contrasting environments is rarely considered. In this paper we describe an experiment in which we compare the P-nutrition of winter and spring barley genotypes from an association genetic-mapping population grown in a field trial with different cultivation treatments (conventional plough vs. minimum tillage) which had been established over a number of years. We demonstrate that, while there is significant variation between genotypes in their P nutrition, this variation is not comparable between cultivation treatments and only one winter barley genotype (cv. Gleam) has beneficial P-use efficiency traits in both cultivation systems. Analysis of the association genetic-mapping population demonstrated that there was a strong environmental component in the genotypic variation, with more significant associations of shoot P concentration with known SNP (Single Nucleotide Polymorphism) markers when the population was grown in minimum tillage treatments. These data suggest that it may be possible to identify genetic components to variation in P nutrition in barley, but that a large interaction with environmental variables may limit the usefulness of any genes or markers discovered for improving P-use efficiency to the conditions under which the screening was performed.     

Banksia species (Proteaceae) from severely phosphorus-impoverished soils exhibit extreme efficiency in the use and re-mobilization of phosphorus

Banksia species (Proteaceae) occur on some of the most phosphorus (P)-impoverished soils in the world. We hypothesized that Banksia spp. maximize P-use efficiency through high photosynthetic P-use efficiency, long leaf lifespan (P residence time), effective P re-mobilization from senescing leaves, and maximizing seed P concentration. Field and glasshouse experiments were conducted to quantify P-use efficiency in nine Banksia species. Leaf P concentrations for all species were extremely low (0.14-0.32 mg P g(-1) DM) compared with leaf P in other species reported and low relative to other plant nutrients in Banksia spp.; however, moderately high rates of photosynthesis (13.8-21.7 micromol CO2 m(-2) s(-1)), were measured. Some of the Banksia spp. had greater P proficiency (i.e. final P concentration in senesced leaves after re-mobilization; range: 27-196 microg P g(-1) DM) than values reported for any other species in the literature. Seeds exhibited significantly higher P concentrations (6.6-12.2 mg P g(-1 )DM) than leaves, and species that sprout after fire ('re-sprouters') had significantly greater seed mass and P content than species that are killed by fire and regenerate from seed ('seeders'). Seeds contained only small amounts of polyphosphate (between 1.3 and 6 microg g(-1) DM), and this was not correlated with P concentration or fire response. Based on the evidence in the present study, we conclude that Banksia species are highly efficient in their use of P, explaining, in part, their success on P-impoverished soils, with little variation between species.     

氮磷添加对巨桉幼苗生物量分配和C:N:P化学计量特征的影响

巨桉(Eucalyptus grandis)是一种优良的速生用材树种, 了解氮(N)和磷(P)对巨桉生长、养分限制、化学计量特征的影响对于科学合理施肥具有重要意义。该实验以巨桉无性系组培苗为研究对象, 通过在酸性紫色土中设置不同施N或施P梯度, 研究巨桉幼苗各器官(根、茎、叶)生物量及碳(C)、N、P的分配和化学计量特征以及巨桉生长的养分限制状况。结果表明: 施N处理对巨桉根茎叶及总生物量的影响极显著, 增加了地上部分的生物量比例而显著降低了根系的生物量比例; 施P对巨桉幼苗总生物量影响不显著, 但显著提高了根的生物量分配比例, 对茎和叶的生物量分配没有显著影响。施N或施P显著改变了巨桉幼苗的N、P含量和化学计量比, 同时也显著影响了土壤与植物N:P的关系。施N可以促使酸性紫色土条件下巨桉对N的吸收而抑制对P的吸收, 施P则促进巨桉幼苗对P的吸收。施N对巨桉幼苗根茎叶的C、N、P分配特征有极显著影响, 而施P对巨桉幼苗根茎叶的C、N、P分配没有显著影响。施N极显著降低了巨桉幼苗N的利用率, 显著提高了P的利用率, 而施P处理极显著降低了巨桉幼苗P的利用率。从巨桉生物量沿施肥梯度和N:P的变化规律可以判断, 当叶片N:P < 15时, 巨桉的生长主要受到N的限制作用。施N可以显著地提高根茎叶的N:P比值, 缓解巨桉缺N的现象, 施P则进一步加剧了N元素的缺乏。     

Mapping of QTLs for phosphorus-deficiency tolerance in rice (Oryza sativa L.)

 Phosphorus (P) deficiency of soils is a major yield-limiting factor in rice production. Increasing the P-deficiency tolerance of rice cultivars may represent a more cost-effective solution than relying on fertilizer application. The objective of this study was to identify putative QTLs for P-deficiency tolerance in rice, using 98 backcross inbred lines derived from a japonica×indica cross and genotyped at 245 RFLP marker loci. Lines were grown on P-deficient soil and P uptake, internal P-use efficiency, dry weight, and tiller number were determined. Three QTLs were identified for dry weight and four QTLs for P uptake, together explaining 45.4% and 54.5% of the variation for the respective traits. Peaks for both traits were in good agreement which was to be expected considering the tight correlation of r=0.96 between dry weight and P uptake. For both traits the QTL linked to marker C443 on chromosome 12 had a major effect. Two of the three QTLs detected for internal P-use efficiency, including the major one on chromosome 12, coincided with QTLs for P uptake; however, whereas indica alleles increased P uptake they reduced P-use efficiency. We concluded that this was not due to the tight linkage of two genes in repulsion but rather due to an indirect effect of P uptake on P-use efficiency. Most lines with high use efficiency were characterized by very low P uptake and dry weight and apparently experienced extreme P-deficiency stress. Their higher P-use efficiency was thus the result of highly sub-optimal tissue-P concentrations and did not represent a positive adaptation to low P availability. The number of tillers produced under P deficiency is viewed as an indirect indicator of P-deficiency tolerance in rice. In addition to the major QTL on chromosome 12 already identified for all other traits, two QTLs on chromosome 4 and 12 were identified for tiller number. Their position, however, coincided with QTLs for tiller number reported elsewhere under P-sufficient conditions and therefore appear to be not related to P-deficiency tolerance. In this study P-deficiency tolerance was mainly caused by differences in P uptake and not in P-use efficiency. Using a trait indirectly related to P-deficiency tolerance such as tiller number, we detected a major QTL but none of the minor QTLs detected for P uptake or dry weight. Received: 9 February 1998 / Accepted: 29 April 1998     

Rice cultivar evaluation for phosphorus use efficiency

Phosphorus deficiency is one of the most growth-limiting factors in acid soils in various parts of the world. The objective of this study was to screen 25 rice cultivars (Oryza sativa L.) at low, medium, and high levels of soil P. Number of tillers, root length, plant height, root dry weight and shoot dry weight were related to tissue P concentrations, P uptake and P-use efficiency. Shoot weight was found to be the plant parameter most sensitive to P deficiency. Significant cultivar differences in P use efficiency were found. Phosphorus use efficiency was higher in roots than shoots and decreased with increasing levels of soil P. Positive correlations were found among growth parameters such as plant height, tillers, root and shoot weight, and P content of roots and shoots. These results indicate selection of rice cultivars for satisfactory performance under low P availability can be carried out using shoot and root dry weight as criteria.     

Transport and partitioning of phosphorus in wheat as affected by P withdrawal during flag-leaf expansion

Increasing P-use efficiency within the plant is one of the acclimations to P-limiting conditions. In this work, we studied the effects of P withdrawal during flag-leaf expansion on sink-source relationships and P-use efficiency in two detillered wheat cultivars (Triticum aestivum L., CA9325 and JM2) under controlled conditions. The study period was divided into two phases of one month each. In the first period after withdrawing P from the medium, the rates of dry weight gain were unaffected compared with the control plant. However, the net dry matter deposition in the ear, and P remobilization within the plant were accelerated in both cultivars. In control plants and in the first period, P transported in the xylem came mainly from the roots current uptake in both cultivars; in the second period, however, phloem retranslocation of P from the shoot and cycling through the root contributed 86% in CA9325 and 95% in JM2 to the xylem-transported P. In the P-deficient plants of both cultivars, almost all of the P transported in the xylem was remobilized, exported from vegetative organs and recycled through the phloem. Over the entire duration of the experiment, the net dry matter deposition and P allocation to grains were not synchronous, indicating independent regulatory processes. Although withdrawing P from the medium markedly reduced the net dry weight gain of whole plants in both cultivars, the final dry weight of the grains was hardly influenced. The percentage of grain dry weight to whole plant dry weight increased from 42.5% in control plants to 44.7% in P-deficient plants in CA9325, and from 41.0% to 45.0% in JM2, and that of P increased from 24.8% to 87.7% and from 25.5% to 84.3%, respectively. The results showed that withdrawing P from the medium during flag-leaf expansion did not influence grain growth and its final P content. The possible mechanisms to regulate P redistribution and reutilization in plants are discussed.     

Plant and microbial strategies to improve the phosphorus efficiency of agriculture

Background

Agricultural production is often limited by low phosphorus (P) availability. In developing countries, which have limited access to P fertiliser, there is a need to develop plants that are more efficient at low soil P. In fertilised and intensive systems, P-efficient plants are required to minimise inefficient use of P-inputs and to reduce potential for loss of P to the environment.

Scope

Three strategies by which plants and microorganisms may improve P-use efficiency are outlined: (i) Root-foraging strategies that improve P acquisition by lowering the critical P requirement of plant growth and allowing agriculture to operate at lower levels of soil P; (ii) P-mining strategies to enhance the desorption, solubilisation or mineralisation of P from sparingly-available sources in soil using root exudates (organic anions, phosphatases), and (iii) improving internal P-utilisation efficiency through the use of plants that yield more per unit of P uptake.

Conclusions

We critically review evidence that more P-efficient plants can be developed by modifying root growth and architecture, through manipulation of root exudates or by managing plant-microbial associations such as arbuscular mycorrhizal fungi and microbial inoculants. Opportunities to develop P-efficient plants through breeding or genetic modification are described and issues that may limit success including potential trade-offs and trait interactions are discussed. Whilst demonstrable progress has been made by selecting plants for root morphological traits, the potential for manipulating root physiological traits or selecting plants for low internal P concentration has yet to be realised.     

磷的生物有效性对山地生态系统的影响

磷(P)的生物有效性对山地生态系统的发育和稳定至关重要。由于大气CO_2浓度升高和N沉降增加,生态系统C、N和P的化学计量比失衡,P的生物有效性受到更多关注。近年来山地系统中P的研究不断深入,2004—2013年间ISI Web of Knowledge中相关研究论文几乎是此前近百年的3倍。总结了山地生态系统中P的生物有效性的特点及其对植物物种多样性和初级生产力的影响。山地生态系统P的生物有效性因垂直高差和地形梯度空间变异明显,快速物质运移和生物过程是控制山地生态系统P的生物有效性的关键因素。P的生物有效性可以影响山地生态系统物种多样性和初级生产力,其对初级生产力的限制存在于全球范围内的山地生态系统。当P的生物有效性发生改变时,山地生态系统的结构越复杂,其植物物种多样性和初级生产力的响应可能会越平缓。全球变化的重要驱动因子(如增温和N沉降增加)可以直接或间接地改变山地生态系统P的生物有效性,因此需要在山地生态系统中加强长期监测和养分控制实验,并结合新型P分析技术,以期认识山地生态系统P的生物有效性的现状、变化趋势和对生态系统的影响,从而为适应全球变化背景下山地生态系统养分状况的改变提供依据。     

Microbial consortium increases maize productivity and reduces grain phosphorus concentration under field conditions

BackgroundThe use of microbes that improve plant phosphorus (P) use efficiency is an avenue to boost crop yields while alleviating environmental impacts. We tested three microbial inoculants (Rhizoglomus irregulare alone – designated AMF; Pseudomonas putida alone – designated PSB; and R. irregulare and P. putida in consortium – designated AMF+PSB), combined with chemical fertilizers, in an intensive maize agricultural system.ResultsAs hypothesized: (i) despite the native soil microbial community and the application of P fertilizer, the microbial inoculants enhanced plant P uptake from the soil by 14–60%, and consequently improved P acquisition efficiency; (ii) PSB and AMF+PSB plants produced ±50% more biomass per unit of P taken up, and consequently enhanced plant internal P use efficiency (i.e. the biomass produced per unit of P); and (iii) the combined inoculation of AMF and PSB provided the best results in terms of productivity and P use efficiency. Further, the microbial inoculants altered P allocation within the plant, reducing grain P concentration.ConclusionBy testing the microbial inoculants under field conditions, our study clearly shows that the microbial consortium (AMF+PSB) increased maize productivity, and at the same time improved P use efficiency. Further, the use of these microbial inoculants was shown to be compatible with conventional agricultural management practices.     

Influence of temperature and soil nitrogen and phosphorus availabilities on fine-root productivity in tropical rainforests on Mount Kinabalu,Borneo

We investigated how temperature and nutrient availability regulate fine-root productivity in nine tropical rainforest ecosystems on two altitudinal gradients with contrasting soil phosphorus (P) availabilities on Mount Kinabalu, Borneo. We measured the productivity and the nutrient contents of fine roots, and analyzed the relationships between fine-root parameters and environmental factors. The fine-root net primary productivity (NPP), total NPP, and ratio of fine-root NPP to total NPP differed greatly among the sites, ranging from 72 to 228 (g m^?2 year^?1), 281–2240 (g m^?2 year^?1), and 0.06–0.30, respectively. A multiple-regression analysis suggested a positive effect of P availability on total NPP, whereas fine-root NPP was positively correlated with mean annual temperature and with P and negatively correlated with N. The biomass and longevity of fine roots increased in response to the impoverishment of soil P. The carbon (C) to P ratio (C/P) of fine roots was significantly and positively correlated with the P-use efficiency of above-ground litter production, indicating that tropical rainforest trees dilute P in fine roots to maintain the C allocation ratio to these roots. We highlighted the mechanisms regulating the fine-root productivity of tropical rainforest ecosystems in relation to the magnitude of nutrient deficiency. The trees showed C-conservation mechanisms rather than C investment as responses to decreasing soil P availability, which demonstrates that the below-ground systems at these sites are strongly limited by P, similar to the above-ground systems.     

Plant biotechnology for lignocellulosic biofuel production

Lignocelluloses from plant cell walls are attractive resources for sustainable biofuel production. However, conversion of lignocellulose to biofuel is more expensive than other current technologies, due to the costs of chemical pretreatment and enzyme hydrolysis for cell wall deconstruction. Recalcitrance of cell walls to deconstruction has been reduced in many plant species by modifying plant cell walls through biotechnology. These results have been achieved by reducing lignin content and altering its composition and structure. Reduction of recalcitrance has also been achieved by manipulating hemicellulose biosynthesis and by overexpression of bacterial enzymes in plants to disrupt linkages in the lignin–carbohydrate complexes. These modified plants often have improved saccharification yield and higher ethanol production. Cell wall‐degrading (CWD) enzymes from bacteria and fungi have been expressed at high levels in plants to increase the efficiency of saccharification compared with exogenous addition of cellulolytic enzymes. In planta expression of heat‐stable CWD enzymes from bacterial thermophiles has made autohydrolysis possible. Transgenic plants can be engineered to reduce recalcitrance without any yield penalty, indicating that successful cell wall modification can be achieved without impacting cell wall integrity or plant development. A more complete understanding of cell wall formation and structure should greatly improve lignocellulosic feedstocks and reduce the cost of biofuel production.     

Genetic diversity for mycorrhizal symbiosis and phosphate transporters in rice

Phosphorus(P) is a major plant nutrient and developing crops with higher P-use efficiency is an important breeding goal.In this context we have conducted a comparative study of irrigated and rainfed rice varieties to assess genotypic differences in colonization with arbuscular mycorrhizal(AM) fungi and expression of different P transporter genes.Plants were grown in three different soil samples from a rice farm in the Philippines.The data show that AM symbiosis in all varieties was established after 4 weeks of growth under aerobic conditions and that,in soil derived from a rice paddy,natural AM populations recovered within6 weeks.The analysis of AM marker genes(AM1,AM3,AM14) and P transporter genes for the direct Pi uptake(PT2,PT6) and AM-mediated pathway(PT11,PT13) were largely in agreement with the observed root AM colonization providing a useful tool for diversity studies.Interestingly,delayed AM colonization was observed in the aus-type rice varieties which might be due to their different root structure and might confer an advantage for weed competition in the field.The data further showed that P-starvation induced root growth and expression of the high-affinity P transporter PT6 was highest in the irrigated variety IR66 which also maintained grain yield under P-deficient field conditions.     

C4 plants as biofuel feedstocks: optimising biomass production and feedstock quality from a lignocellulosic perspective

The main feedstocks for bioethanol are sugarcane (Saccharum officinarum) and maize (Zea mays), both of which are C(4) grasses, highly efficient at converting solar energy into chemical energy, and both are food crops. As the systems for lignocellulosic bioethanol production become more efficient and cost effective, plant biomass from any source may be used as a feedstock for bioethanol production. Thus, a move away from using food plants to make fuel is possible, and sources of biomass such as wood from forestry and plant waste from cropping may be used. However, the bioethanol industry will need a continuous and reliable supply of biomass that can be produced at a low cost and with minimal use of water, fertilizer and arable land. As many C(4) plants have high light, water and nitrogen use efficiency, as compared with C(3) species, they are ideal as feedstock crops. We consider the productivity and resource use of a number of candidate plant species, and discuss biomass 'quality', that is, the composition of the plant cell wall.     

A damped precipitation‐driven,bottom‐up model for deer mouse population abundance in the northwestern United States

Small‐mammal population densities can be regulated by bottom‐up (food availability) and top‐down (predation) forces. In 1993, an El Niño Southern Oscillation event was followed by a cluster of human hantavirus with pulmonary syndrome in the southwestern United States. An upward trophic cascade hypothesis was proposed as an explanation for the outbreak: Increased plant productivity as a consequence of El Niño precipitations led to an unusual increase in distribution and abundance of deer mice (Peromyscus maniculatus ; reservoir host of Sin Nombre virus). Could such drastic events occur in mesic habitats, where plant productivity in response to climate conditions is likely to be much less dramatic? In this work, we investigate to what extent deer mouse populations follow a precipitation‐driven, bottom‐up model in central and western Montana and discuss important conditions for such a model to be possible. We found positive correlations between deer mouse abundance and on‐the‐ground measured plant productivity with a several‐month lag in three of six study sites. This effect was weaker when deer mouse populations were more abundant, indicating density‐dependent effects. Dispersal resulting from territoriality may be important in attenuating local density increments in spite of high food availability. In addition, there is evidence that population abundance in the study area could respond to other abiotic factors. In particular, precipitation in the form of snow may reduce deer mice survival, thus compensating the benefits of improved plant productivity. Deer mouse populations in Montana study sites follow complex dynamics determined by multiple limiting factors, leading to a damped precipitation‐driven bottom‐up regulation. This prevents dramatic changes in rodent abundances after sudden increments of food availability, such as those observed in other regions.     

Plant nutrient efficiency: A comparison of definitions and suggested improvement

Selection of plant cultivars tolerant of low nutrient supply may increase productivity on low fertility soils and reduce fertilizer requirements. Considerable effort has been directed towards identifying nutrient efficient species and germplasms, but the many different definitions for efficiency make the use of the term ambiguous. The concept of nutrient efficiency was evaluated using data from a study of differences in germplasm response to phosphorus (P) availability in white clover (Trifolium repens L.) and alfalfa (Medicago sativa L.) grown in a sand-alumina culture. Application of various criteria identified in the literature as measures of nutrient efficiency did not clarify differences between purportedly P efficient and inefficient germplasms. Germplasms differed in maximum shoot and total dry mass and in solution P concentration required to achieve 80% maximum yield, but not in tissue P concentration, internal P utilization, or P uptake per unit of fine root dry mass. Differences may have resulted from factors other than efficient use of available P. To reduce the confounding effects that other factors have on nutrient efficiency, we propose that equivalent yields of germplasms be demonstrated where nutrients are not limiting. Mechanisms that enable enhanced nutrient efficiency can be identified less ambiguously using this improved approach.Joint contribution of the Minnesota Agricultural Experimental Station, USDA-ARS and US Dairy Forage Research Center (Minnesota cluster). Paper No. 20,432 of the Minnesota Science Journal Series.Joint contribution of the Minnesota Agricultural Experimental Station, USDA-ARS and US Dairy Forage Research Center (Minnesota cluster). Paper No. 20,432 of the Minnesota Science Journal Series.     

Reconsidering diversity–productivity relationships: directness of productivity estimates matters

Despite extensive research efforts, the controversy over diversity–productivity (D–P) patterns in natural communities still looms large. Recent meta‐analyses suggest that unimodal D–P relationships tend to pre‐dominate in plant studies, while positively linear relationships are more common in animal studies. These patterns, however, are based on studies in which productivity is estimated either directly, based on the biomass or energy of the studied organisms, or indirectly, according to the productivity of lower trophic levels, and various surrogates. Our analysis shows that the distribution of D–P patterns is sensitive to the directness of productivity estimates in animal studies but not in plant studies. Analysis of D–P patterns should be based on direct productivity estimates of the studied organisms, especially in comparative meta‐analyses of communities from multiple trophic levels, where productivity is often affected nonlinearly by indirect factors or when complex feedback interactions are expected between productivity and diversity.     

Context dependent effects of ectomycorrhizal species richness on tree seedling productivity

While there has been much recent interest about the relationships between plant diversity and plant productivity, much remains unknown about how the diversity of mycorrhizal fungi affects plant productivity. We investigated the effects of ectomycorrhizal fungal community composition and diversity on the productivity and growth characteristics of seedlings of two tree species ( Pinus sylvetris and Betula pendula ) as well as their interactions with each other. This involved setting up a mycorrhizal fungal diversity gradient from one to eight species using a design previously demonstrated to be able to separate diversity effects from compositional effects. We found that the eight mycorrhizal fungal species differed in their effects on seedling productivity and that the nature of effects was determined by the fertility of the substrate. Fungal species richness effects were also important in affecting seedling productivity over and above what could be explained by "sampling effect" but only in some situations. For B. pendula in a low fertility substrate there were clear positive causative effects between fungal species richness and productivity with the eight species treatment having over double the productivity of any of the eight monoculture treatments; no diversity effects were, however, detected in a high fertility substrate. For P. sylvestris in a high fertility substrate there were significant negative effects of fungal diversity on productivity while in a low fertility substrate no effects were apparent. The possible mechanistic bases for these results are discussed. The growth of P. sylvestris relative to that of B. pendula when grown in combination was unaffected by mycorrhizal treatments. Our results provide clear evidence that effects of mycorrhizal fungal diversity on productivity are context dependent and may be positive, negative or neutral depending on the situation considered.     

Nutrient transport in mycorrhizas: structure,physiology and consequences for efficiency of the symbiosis

Nutrient transport in mycorrhizas occurs across specialized interfaces which are the result of corrdinated development of the organisms. The structural modifications give rise to large areas of either inter- or intra-cellular interface in which wall synthesis is frequently modified and in which altered distribution of membrane bound ATPases is important, particularly with respect to mechanisms that may be involved in bidirectional transfer of nutrients. Except in orchid mycorrhizas, net movement of organic carbon from plant to fungus occurs, complemented by mineral nutrient movement in the opposite direction. The general consensus is that sustained transfer at rates that will maintain the growth and development of the organisms requires increases in the rates at which nutrients are lost from the organisms; possible mechanisms for this are discussed. The transfer processes are essential in determining both plant and fungal productivity and an approach to calculating the efficiency of the symbiosis in terms of the expenditure of carbon (or of phosphorus) is discussed.     

Ecophysiological response in leaves of Caragana microphylla to different soil phosphorus levels

Phosphorus (P) is one of the limiting mineral nutrient elements in the typical steppe of Inner Mongolia, China. In order to find out the adaptive strategy of Caragana microphylla to low soil P status, we grew plants in P-deficient soil in April 2009 and gave a gradient of P addition ranging from 0 to 60 mg(P) kg^?1(soil) from May 2010. Leaf traits were measured in September 2010. Both leaf growth and light-saturated photosynthetic rate (P _max) were similar among different groups. Leaf nitrogen (N):P ratio indicated that the growth of C. microphylla was not P-limited in most of the Inner Mongolia typical steppe, which had an average soil available P content equal to 3.61 mg kg^?1. The optimal P addition was 20 mg(P) kg^?1(soil) for two-year-old plants of C. microphylla. Leaf mass area (LMA) and leaf dry matter content (LDMC) were enhanced with low P, and significantly negatively correlated with photosynthetic N-use efficiency (PNUE). Photosynthetic P-use efficiency (PPUE) increased with decreasing soil P and increasing leaf inorganic P (Pi): organic P (Po) ratio, and showed no significant negative correlation with LMA or LDMC. P _max of C. microphylla did not decline so sharply as it was anticipated. The reason for this phenomenon might be due to the increased PPUE through regulating the leaf total P allocation. C. microphylla had high P-use efficiency via both high PPUE and long P-retention time at low-P supply. The adaptation of C. microphylla to low-P supply provided a new explanation for the increased distribution of the species in the degraded natural grassland in Inner Mongolia, China.