Green and senescent leaf phenolics showed spatial autocorrelation in a Quercus robur population in northwestern Spain

Intraspecific variation in polyphenols may be important both for the resistance of plants to herbivory and for nutrient cycling in terrestrial ecosystems. The spatial pattern and scale of polyphenol concentration in natural populations are practically unknown, despite multiple evidence that resources influencing leaf phenolics (such as light or nutrient availability) show spatial dependence at the scale of meters to tens of meters. By using geostatistical methods, we evaluated the spatial variability in polyphenol concentration in green and senescent leaves in a pedunculate oak (Quercus robur L.) population of 125 individuals. The spatial pattern of light and nutrient availability for plants was also evaluated. Leaf polyphenols were more variable than other leaf properties, and green polyphenols were significantly correlated with radiation, leaf mass per unit area and leaf N concentration. Both green and senescent leaf polyphenol concentration showed spatial dependence at distances below 6 to 10 m, with structural (spatial) variance explaining between 50 and 80% of the total variance. The variance not explained by space was assumed to be due to individual genotypic variability. Stochastic maps of leaf phenolics in the area showed that the probability of finding high phenolics level was not random. These findings may be important to understand the spatial heterogeneity of plant-herbivore interactions, leaf litter decomposition and mineralization rate.     

Effects of artificial grassland establishment on soil nutrients and carbon properties in a black-soil-type degraded grassland

Despite a growing knowledge of nutrient limitation for mangrove species and how mangroves adapt to low nutrients, there is scant information about the relative importance of N:P ratio and leaf phenolics variability in determining nutrient conservation. In this study, we evaluated possible nutrient conservation strategies of a mangrove Rhizophora stylosa under nutrient limitation. 1. The leaf nutrient concentrations of R. stylosa changed with season, with the highest N concentration in winter and the highest P concentration in spring for both mature and senescent leaves. Leaf N and P concentrations decreased significantly during leaf senescence. Based on N:P ratios R. stylosa forest was N-limited. Accordingly, the nitrogen resorption efficiency (NRE) was significantly higher than phosphorus resorption efficiency (PRE) for the R. stylosa leaves during leaf senescence. The NRE and PRE both reached the highest in the autumn. Average N and P concentrations in the senescent leaves were 0.15% and 0.06% for R. stylosa, respectively, indicating a complete resorption of N and an incomplete resorption of P. There was a significant negative correlation between nitrogen resorption proficiency (NRP) and NRE, meanwhile phosphorus resorption proficiency (PRP) and PRE correlation was also highly significantly. 2. R. stylosa leaves contained relatively high tannin level. Total phenolics, extractable condensed tannins and total condensed tannins contents increased during leaf senescence, and changed between seasons. The lowest concentrations of total phenolics, extractable condensed tannins and total condensed tannins occurred in summer, total phenolics concentrations were inversely related to nitrogen or phosphorus concentrations. 3. Our results confirmed that resorption efficiency during leaf senescence depends on the type of nutrient limitation, and NRE was much higher than PRE under N-limited conditions. R. stylosa forest developed several nutrient conservation strategies in the intertidal coastline surroundings, including high nitrogen resorption efficiency, low nutrient losses and high tannins level.     

Nutrient resorption in tropical savanna forests and woodlands of central Brazil

Nutrient limitation in Brazilian savanna (known as cerrado) presumably causes trees to maximize nutrient resorption from senesced leaves to reduce their dependence on nutrient availability. To assess patterns between nutrient resorption and soil fertility, we measured community-level nitrogen (N), phosphorus (P), and potassium (K) concentrations in mature and senesced leaves and soil fertility in the upper 50 cm soil layer in structurally diverse cerrado ecosystems in the Cuiaba Basin (CB) and Pantanal (PAN) of Mato Grosso, Brazil. Foliar nutrient concentration data were used to estimate resorption efficiency and proficiency, and correlation was used to determine whether resorption efficiency and proficiency varied across soil fertility gradients. We found that N and P resorption proficiency (NRP and PRP, respectively) and P resorption efficiency (PRE) increased significantly as total soil N (NRP) and extractable P (PRP and PRE) declined. In contrast, K resorption efficiency (KRE) declined as soil sand content and bulk density increased, which was likely due to a reduction in soil water-holding capacity. Leaf N/P ratios indicate potential N limitation and/or N + P co-limitation for ecosystems in the PAN and P limitation and/or N + P co-limitation for ecosystems in the CB, while trends in leaf N/K ratios indicate possible K or K + P co-limitation for the CB only. Our results illustrate that cerrado forests and woodlands have highly variable nutrient resorption capacities that vary predictably across soil fertility or textural gradients and indicate that cerrado communities have flexible nutrient resorption that can reduce their dependence on soil nutrient availability.     

元谋干热河谷燥红土和变性土上植物叶片的元素含量及其重吸收效率

采用盆栽试验,研究元谋干热河谷燥红土和变性土上生长的植物叶片以及凋落叶营养元素含量,并分析养分重吸收效率对土壤类型与物种互作的响应.结果表明: 土壤类型对叶片N、P、Ca、Mg、Cu、Zn、Fe、N∶P以及凋落叶N、P、Mn、N∶P均有显著影响;燥红土植物叶片与凋落叶N、Mn含量和N∶P显著高于变性土,而燥红土植物叶片P、Ca、Mg、Fe、Cu、Zn和凋落叶P含量显著低于变性土.燥红土植物叶片N含量较变性土高34.8%,而P含量低40.0%;在叶片凋落时,N、P、K表现为重吸收,而其他元素呈富集状态.燥红土凋落叶Ca、Mg、Mn富集系数显著高于变性土.物种仅对叶片N含量有显著影响,物种与土壤交互作用对植物叶片和凋落叶元素含量影响不显著,表明各土壤类型对不同物种元素含量的影响方式较为一致.土壤类型对植物元素含量的影响可进一步作用于干热河谷植物凋落物分解、植物-土壤的养分反馈以及生物地球化学循环.     

Foliar nutrient resorption patterns of four functional plants along a precipitation gradient on the Tibetan Changtang Plateau

Nutrient resorption from senesced leaves as a nutrient conservation strategy is important for plants to adapt to nutrient deficiency, particularly in alpine and arid environment. However, the leaf nutrient resorption patterns of different functional plants across environmental gradient remain unclear. In this study, we conducted a transect survey of 12 communities to address foliar nitrogen (N) and phosphorus (P) resorption strategies of four functional groups along an eastward increasing precipitation gradient in northern Tibetan Changtang Plateau. Soil nutrient availability, leaf nutrient concentration, and N:P ratio in green leaves ([N:P]_g) were linearly correlated with precipitation. Nitrogen resorption efficiency decreased, whereas phosphorus resorption efficiency except for sedge increased with increasing precipitation, indicating a greater nutrient conservation in nutrient‐poor environment. The surveyed alpine plants except for legume had obviously higher N and P resorption efficiencies than the world mean levels. Legumes had higher N concentrations in green and senesced leaves, but lowest resorption efficiency than nonlegumes. Sedge species had much lower P concentration in senesced leaves but highest P resorption efficiency, suggesting highly competitive P conservation. Leaf nutrient resorption efficiencies of N and P were largely controlled by soil and plant nutrient, and indirectly regulated by precipitation. Nutrient resorption efficiencies were more determined by soil nutrient availability, while resorption proficiencies were more controlled by leaf nutrient and N:P of green leaves. Overall, our results suggest strong internal nutrient cycling through foliar nutrient resorption in the alpine nutrient‐poor ecosystems on the Plateau. The patterns of soil nutrient availability and resorption also imply a transit from more N limitation in the west to a more P limitation in the east Changtang. Our findings offer insights into understanding nutrient conservation strategy in the precipitation and its derived soil nutrient availability gradient.     

Litter decomposition of woody species in shrublands of NW Patagonia: how much do functional groups and microsite conditions influence decomposition?

The study examined the effects of leaf traits, soil microsite, and microclimate characteristics on litter decomposition of the dominant species in two functional groups (FG), deciduous and evergreen, in shrublands in NW Patagonia, Argentina. Leaf traits considered were nutrient concentration (C, N, P, C/N, and N/P) and physical characteristics (area, strength, specific leaf area, and dry matter content). Soil microsite characteristics measured were pH, C, N, P, C/N and water retention capacity, while soil microclimate characteristics recorded were soil and air, temperature and moisture, and solar radiation. Five evergreen and five deciduous woody shrub species were selected. During 1 year, litter and microsite properties were measured below canopy: (i) senescent leaf chemical and physical properties, and the quantity as well as field decomposition of litter and (ii) soil chemistry, and soil and air physical properties. The factors controlling litter decomposition were different for each FG. In deciduous species, C/N ratio had a negative effect on decomposition. In evergreen species, decomposition was affected negatively by leaf carbon and dry matter content. Litter decomposition depended exclusively on the inherent senescent leaves traits. The common decomposition pattern between species of both FG could be attributed to similar leaf traits and the correlation between variables that control decomposition in both groups. Plant nutrient inputs associated with the litter decomposition process did not explain the soil nutrient content. These results suggest that other organic matter sources (roots, branches, and fruits) are more important than leaves on soil fertility.     

黄土高原小流域土壤养分的空间异质性

利用地理信息系统的空间分析功能,通过地统计学的半变异函数定量研究了黄土高原典型小流域土壤养分的空间异质性特征。结果表明;土壤有机质,全氮,有效氮,全磷和有效磷的理论模型均为球状模型,由随机因素引起的空间变异占空间总变异的比例小,其值分别为13.333%,10.938%,22.000%,9.091%和27.536%,反映5种养分具有较强的空间自相关格局,但它们的空间自相关范围具有明显的差异。土壤全氮和有效氮变程小,分别是90m和110m,有机质次之,变程是120m,而土壤全磷和有效磷的变程最大为160m,研究成果将有效地指导土壤的取样设计,以及进行土壤养分的空间内插和制图。     

水分对敦煌阳关湿地芦苇叶片与土壤C、N、P生态化学计量特征的影响

土壤水分是影响干旱区植物养分吸收和利用策略的关键因子之一。研究不同水分梯度叶片与土壤生态化学计量特征,有助于揭示植物对环境变化的响应特征及生态适应性。通过野外调查与实验分析,对敦煌阳关不同水分梯度芦苇叶片与土壤碳(C)、氮(N)、磷(P)生态化学计量特征及其关系进行了研究。结果表明:(1)随土壤含水率升高,叶片C、N、P含量降低,叶片C/N、C/P、N/P升高。(2)随土壤含水率升高,土壤有机碳(OC)、总氮(TN)、总磷(TP)含量及土壤N/P升高,土壤C/N降低,土壤C/P先升后降。(3)低水分梯度叶片N、C/N与土壤N、C/N显著负相关(P0.05),叶片C、P、C/P、N/P与土壤C、P、C/P、N/P无显著相关性(P0.05);高、中水分梯度叶片C、N、P与土壤C、N、P化学计量特征相关性均不显著(P0.05)。低水分梯度叶片受干旱胁迫和土壤养分制约,且能够保持较高的叶养分含量,体现了干旱区湿地植物异质生境下独特的养分调节机制。     

Leaf nutrient concentration as an indicator of Populus and Tamarix response to flooding

Plants growing in infertile environments are able to produce more biomass per unit of nutrient taken up than plants of fertile habitats, and also to minimize nutrients loss by resorbing them from senescing leaves. The leaf nutrient concentration variability of two co-existing riparian tree genera (Populus and Tamarix) along a flood inundation gradient was examined to infer nutrient limitation and to compare nutrient use strategies in the two genera. To that end, seasonal and spatial variability in leaf nitrogen (N) and phosphorus (P) concentration (i.e., % dry mass of N and P) were analyzed in 720 samples of leaves (2 tree genera × 3 seasons × 12 sites × 10 tree replicates). Both Populus and Tamarix showed strong seasonal variability in leaf N and P concentrations, with values decreasing throughout the growing season. However, while N:P atomic ratio remained seasonally constant in Populus (N:P = 33), Tamarix shifted from N:P = 29 in spring to N:P = 36 and 37 in summer and fall. %N, %P and N:P atomic ratios were also spatially variable, but leaf litter N and P concentration (i.e., nutrient resorption proficiency) and leaf litter N:P generally followed the local flood inundation gradient as shown by linear mixed effects models. In particular, nutrient resorption was usually less proficient (higher terminal nutrient concentrations) at higher flood durations (in gravel bars and natural levees), whereas N:P increased in the drier sites (floodplain terrace). At floodplain level, a P-limitation that is higher than N-limitation seems to characterize the plant nutrient circulation in the riparian ecosystem studied. Tamarix was slightly more proficient in P resorption than Populus. The study shows that leaf nutrient concentration (e.g., N and P) derived from nutrient availability is partly controlled by the flood inundation regime and can be used as an indicator of nutrient limitation in forested floodplains. Subtle differences between tree genera provide an additional, novel explanation for the recent expansion of Tamarix in many arid and semi-arid rivers with altered hydrogeomorphic regimes.     

Changes in phosphorus fractions caused by increased microbial activity in forest soil in a short-term incubation study

The effects of adding larch (Larix kaempferi) leaf litter and nitrogen (N) on microbial activity and phosphorus (P) fractions in forest soil were examined in a short-term (28-d) laboratory incubation study. The soil was analyzed using a modified Hedley sequential extraction procedure and an acid phosphatase assay. The addition of larch litter and N increased the acid phosphatase activity and decreased the labile P (H₂O-P + NaHCO₃-P) concentration. Compared with addition of larch litter only, addition of both inputs decreased the proportion of inorganic P (Pi) and increased that of organic P (Po) in the NaOH fraction, bound to aluminum and iron oxides. The results of nutrient (carbon, N, or P) addition indicated that acid phosphatase was synthesized to acquire P. This study suggests that, in this forest soil, P in the H₂O-P + NaHCO₃-P and in the NaOH-Pi fractions was available for soil microorganisms to decompose leaf litter and that increase in microbial activity eventually translated in an increase in the proportion of Po found in the NaOH fraction in this forest soil.     

Coloration and phenology manifest nutrient variability in senesced leaves of 46 temperate deciduous woody species

颜色和物候表明46种温带落叶木本植物衰老叶片的养分变异 不同共生植物的叶片养分含量差异显著,反映了不同的叶片养分利用策略。然而,衰老叶片养分的种间变异及其驱动因素尚不清楚。本研究旨在探讨衰老叶片养分的种间变异及其驱动因素。我们在中国东北的帽儿山森林生态系统研究站测定了46种共存温带落叶木本植物新鲜落叶的碳、氮、磷浓度。 采用随机森林模型量化10个生物因素(菌根类型、固氮类型、生长形态、耐阴性、叶片质地、变色程度、变色类型、叶片变色峰期、落叶峰期和落叶末期)的相对重要性。研究结果表明,落叶氮浓度种间变化为4倍,磷浓度变化达9倍。较高的氮和磷平均浓度(15.38和1.24 mg g⁻¹)表明该森林氮和磷限制较弱。功能群仅对特定养分及其比值有显著影响。磷浓度、氮磷比与外生菌根树种的落叶高峰日和落叶结束日呈负相关。颜色鲜艳的叶片(红色＞棕色＞黄色＞黄绿色＞绿色)倾向于比绿色叶片氮和磷浓度更低而碳氮比和碳磷比较高。随机森林模型表明,秋季叶变色和落叶物候贡献了80%的种间变异解释量。这些结果增加了我们对温带森林木本植物营养策略之衰老叶片养分变异性的理解。     

内蒙古草地叶片磷含量与土壤有效磷的关系

植物体内特别是叶片的P含量特征及其与环境的关系一直是植物生理生态学研究的一个热点。已有研究发现, 与全球植物数据相比, 中国植物叶片的P含量相对较低, 导致N/P高于全球平均水平, 并推测这是由于中国土壤全P含量较低引起的。该研究选取内蒙古草地来验证这一假设, 分析了36个样地57种优势植物叶片的P含量与土壤全P和有效P含量的关系。主要结果如下: 内蒙古草地叶片P含量较低而N/P较高, 与之前的研究结论一致; 在种群、样地和物种3个水平上, 叶片P含量、N/P与土壤P含量都没有显著的相关关系, 尽管土壤有效P含量的解释力高于土壤全P; 另一方面, 内蒙古草地土壤的有效P含量与全国土壤普查的结果接近, 高于美国及澳大利亚的平均值, 但低于世界土壤信息库里报道的全球土壤有效P平均值。鉴于内蒙古草地土壤的全P和有效P含量都不能准确反映叶片P含量, 且土壤的有效P含量也并不明显低于世界其他地区, 因此植物叶片P含量低、N/P高是由于土壤P含量低引起的这一假设在内蒙古草地不成立, 而且叶片P含量也与土壤P的可利用性无关。     

Plasticity in latitudinal patterns of leaf N and P of Oryza rufipogon in China

Characterising the adaptability in nature of plant stoichiometric patterns across geographic or environmental gradients is important in advancing our understanding of the organisation of plant–nutrient relationships. We examined correlations between plant nutrient traits, latitude, longitude, climate and soil variables in 34 populations of Oryza rufipogon across its range. We further compared the responses of population transplants at two experimental gardens: one beyond its northern natural range and another near the southern limit, to assess the nature of geographic variation in plant nutrients. The study showed that leaf P of O. rufipogon in the field was negatively correlated with latitude and largely depended on temperature and soil P availability. Leaf N was not related to latitude but was significantly correlated with precipitation and soil N concentration. Leaf N:P ratio was largely determined by absorption efficiency of P. Transplantation revealed that there were no significant associations of leaf nutrients with geographic, climatic or soil variables of origin in either of the experimental gardens, indicating phenotypic plasticity. However, examination of relationships between response ratios of leaf nutrients and change ratio of climate and soil environments, as well as norms of reaction in the transplantation experiment, revealed more complexity, suggesting both substantial genotypic diversity and the existence of genotype × environment interactions in these populations of O. rufipogon. These data indicate that adaptive plasticity response of plants to temperature and soil P availability significantly explain the observed shifts in leaf N, P and N:P of O. rufipogon along latitudinal gradients.     

Tree species effects on coupled cycles of carbon, nitrogen, and acidity in mineral soils at a common garden experiment

Forest biogeochemical cycles are shaped by effects of dominant tree species on soils, but the underlying mechanisms are not well understood. We investigated effects of temperate tree species on interactions among carbon (C), nitrogen (N), and acidity in mineral soils from an experiment with replicated monocultures of 14 tree species. To identify how trees affected these soil properties, we evaluated correlations among species-level characteristics (e.g. nutrient concentrations in leaf litter, wood, and roots), stand-level properties (e.g. nutrient fluxes through leaf litterfall, nutrient pools in stemwood), and components of soil C, N, and cation cycles. Total extractable acidity (acidity_tot) was correlated positively with mineral soil C stocks (R ²  = 0.72, P < 0.001), such that a nearly two-fold increase in acidity_tot was associated with a more than two-fold increase of organic C. We attribute this correlation to effects of tree species on soil acidification and subsequent mineral weathering reactions, which make hydrolyzing cations available for stabilization of soil organic matter. The effects of tree species on soil acidity were better understood by measuring multiple components of soil acidity, including pH, the abundance of hydrolyzing cations in soil solutions and on cation exchange sites, and acidity_tot. Soil pH and acidity_tot were correlated with proton-producing components of the soil N cycle (e.g. nitrification), which were positively correlated with species-level variability in fine root N concentrations. Soluble components of soil acidity, such as aluminum in saturated paste extracts, were more strongly related to plant traits associated with calcium cycling, including leaf and root calcium concentrations. Our results suggest conceptual models of plant impacts on soil biogeochemistry should be revised to account for underappreciated plant traits and biogeochemical processes.     

Nitrogen fertilization and fire act independently on foliar stoichiometry in a temperate steppe

Nitrogen (N) fertilization, as a grassland management strategy, has been widely used to improve forage quality and increase the productivity of grasslands degraded by overstocking. It is widely accepted that N addition will alter ecosystem structure and function, and that these effects may be altered by natural disturbances, such as fire. We examined the effects of annual burning and N fertilization (17.5 g N m^?2 year^?1, at a surplus rate in order to simulate agriculture treatment) on foliar chemistry and stoichiometric ratios of eight dominant species (Leymus chinensis, Stipa grandis, Cleistogenes squarrosa, Potentilla bifurca, Thalictrum squarrosum, Artemisia frigida, Kochia prostrata and Caragana microphylla) in a temperate steppe in northern China. After 3 years of treatments, annual burning significantly increased soil extractable phosphorus (P) concentration but showed no effects on soil inorganic N concentration, whereas N fertilization caused a significant increase in inorganic N concentration but not of extractable P. Species differed substantially with respect to all nutritional and stoichiometric variables. Both annual burning and N fertilization caused significant increases in foliar N and P concentrations and thus decreases in carbon (C):N and C:P ratios. While annual burning showed no effects on N:P ratios, N fertilization produced higher N:P ratios. However, species responded idiosyncratically to both fire and N fertilization in terms of foliar N concentration, C:N and N:P ratio. In addition, there was no interaction between fire and N fertilization that affected all variables. This study suggests that both annual burning and N fertilization have direct impacts on plant elemental composition and that fire- and N addition-induced changes of community composition may have important consequences for plant-mediated biogeochemical cycling pathways in temperate steppe ecosystem.     

Soil C:N:P stoichiometry responds to vegetation change from grassland to woodland

Woody encroachment has been a major land cover change in dryland ecosystems during the past century. While numerous studies have demonstrated strong effects of woody encroachment on soil carbon (C), nitrogen (N), and phosphorus (P) storage, far less is known about the plasticity of soil C:N:P stoichiometry in response to woody encroachment. We assessed landscape-scale patterns of spatial heterogeneity in soil C:N:P ratios throughout a 1.2 m soil profile in a region where grassland is being replaced by a diverse assemblage of subtropical woody plants dominated by Prosopis glandulosa, an N₂-fixing tree. Woody species had leaf and fine root C:N:P ratios significantly different from grasses. Variation in soil C:N ratios in both horizontal and vertical planes was remarkably smaller than that of soil N:P and C:P ratios. Spatial patterns of soil C:N ratio throughout the profile were not strongly related to vegetation cover. In contrast, spatial patterns of soil N:P and C:P ratios displayed a strong resemblance to that of vegetation cover throughout the soil profile. Within the uppermost soil layer (0–5 cm), soil N:P and C:P ratios were higher underneath woody patches while lower within the grassland; however, this pattern was reversed in subsurface soils (15–120 cm). These results indicate a complex response of soil C:N:P stoichiometry to vegetation change, which could have important implications for understanding C, N, and P interactions and nutrient limitations in dryland ecosystems.     

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。     

Is primary production in holm oak forests nutrient limited?

Correlations between primary production and patterns of nutrient use and nutrient availability were investigated in 18 plots in closed holm oak (Quercus ilex L.) stands in the Montseny mountains (NE Spain), searching for evidence of nutrient limitation on primary production. The plots spanned a range of altitudes and slope aspects within a catchment. Nutrients considered were nitrogen (N), phosphorus (P), potassium (K) and magnesium (Mg) in plant samples, and the above plus calcium (Ca) and sodium (Na) in the soil. Primary production was found by summing the annual aboveground biomass increment to the annual litterfall. Across plots, primary production was correlated with the annual return of nutrients in litterfall, but this relationship probably arose from the common effects of the amount of litterfall on both primary production and nutrient return, and not from any nutrient limitation. Primary production was not significantly correlated with nutrient concentrations in mature leaves nor leaf litterfall, nor with absolute or relative foliar retranslocation of nutrients before leaf abscission, nor with the concentration and content (kg/ha) of total N, extractable P, and exchangeable K, Mg, Ca and Na in the upper mineral soil. We conclude that there is no correlational evidence that primary production is nutrient limited in this holm oak forest.     

Small-scale spatial heterogeneity in the vertical distribution of soil nutrients has limited effects on the growth and development of Prosopis glandulosa seedlings

We examined the above- and below-ground responses of seedlings of the woody shrub Prosopis glandulosa to the spatial heterogeneity of soil nutrients within the root zone. We performed a microcosm experiment where seedlings were grown with different combinations of nutrients [nitrogen (N), phosphorus (P), and both combined (NP)] and under different levels of nutrient heterogeneity (nutrients supplied as patches located in the bottom and/or upper portion of rooting zone versus homogeneous distribution). Seedling morphology and biomass did not show a strong response to changes in nutrient ion or spatial heterogeneity. Height, number of leaves, and specific leaf area did not vary significantly between treatments. The number of leaves, foliar biomass, stem biomass and biomass allocation to stems of seedlings showed more responsiveness to the addition of N and NP than to the addition of P. The spatial heterogeneity of nutrients affected the diameter, root biomass and leaf N content. Seedlings had higher diameter and root biomass when the nutrients were homogeneously distributed as compared to their placement as patches in the bottom of the microcosms. Their leaf N concentration increased in those treatments where the nutrient patch was located in the lower half as compared to the upper half of the microcosms. Root foraging responses to nutrient patches varied with their location. Significant root proliferation was observed when patches of N, P and NP were located in the upper portion of the rooting zone; when they were located in the lower portion such a response was observed only for P. Despite our findings that Prosopis seedlings have a low overall responsiveness to small-scale vertical differences in soil nutrient heterogeneity, our results suggest that these differences may modify the growth dynamics of the secondary roots of this ecologically important dryland species during the early stages of its development.