Leaf mass loss in wetland graminoids during senescence

Mass loss of senescing leaves is an important part of plant biomass turnover and has consequences for assessment of ecosystem productivity, ecosystem nutrient use efficiency, and plant nutrient resorption efficiency. Data, however, on mass loss are scarce, and often based on leaf area as the reference base. This leads to an underestimation of the mass loss, as leaf area itself shrinks during senescence. Furthermore, the few existing studies have almost exclusively used woody species. The purpose of the present study was twofold: i) to assess leaf mass loss during senescence in herbaceous species, with the example of five wetland graminoids and, ii) to compare two different methods of mass loss assessment (two species). Assuming that leaf length does not change during senescence, we assessed leaf mass per leaf length prior to and after senescence. We also estimated pre‐senescence leaf mass nondestructively based on leaf length, width and thickness. For Typha latifolia and Carex stricta, two species with graminoid type leaves but contrasting leaf structure, both methods delivered almost identical results. After the first assessment of leaf mass on July 7th, T. latifolia leaf mass initially increased by 13%, and then decreased to be 12% below the original mass after senescence. C. stricta leaf mass remained stable until senescence, but decreased then by 33%. In a second experiment, the mass of 100 mm pieces of leaves was measured before and after senescence. Calamagrostis canadensis, Carex rostrata and C. stricta lost 23–57% of their leaf mass during senescence, whereas Glyceria canadensis did not show any mass loss. We conclude that mass loss of senescing leaves of herbaceous plants can be considerable and should not be neglected in studies of productivity, nutrient use efficiency or nutrient resorption. For species with no shrinking leaf length during senescence, mass loss can be measured with leaf length as the base whereas for others, pre‐senescent mass can be estimated on the basis of leaf dimensions.     

Current measures of nutrient resorption efficiency lead to a substantial underestimation of real resorption efficiency: facts and solutions

Nutrient resorption is an important process during leaf senescence, which helps plants to minimize nutrient losses. To quantify nutrient resorption, the parameter resorption efficiency is commonly used. This parameter describes the percentage of the nutrient pool withdrawn before leaf abscission. The nutrient pool is generally expressed on the basis of leaf mass or leaf area, assuming that these bases do not change during senescence. In this paper we firstly present a mathematical formula describing the effect of change in measurement basis on the difference between the real resorption efficiency (RRE) value and the measured resorption efficiency (MRE). This formula shows that even moderate senescence-related changes in a measurement basis can lead to considerable underestimation of RRE. Secondly, to estimate the general change in measurement basis we quantified leaf mass loss and leaf shrinkage during senescence from literature data. These data shows that mass loss percentages can be as high as 40%, and leaf shrinkage can be up to 20%. This level of change in basis seriously compromises the MRE when not corrected for. Using our formula and the reported average literature values of changes in leaf mass (21%) and leaf shrinkage (11%) during senescence, we calculated that the average RRE for nitrogen and phosphorous of terrestrial plants is 6% (leaf area) to 10% (leaf mass) higher than the 50%, respectively 52% as reported by Aerts (1996) . This implies that nutrient resorption from senescing leaves is even more important for nutrient retention in terrestrial plants than thought so far. We advocate that preselecting leaves and monitoring the measurement basis throughout the duration of the experiment should minimize the difference between MRE and RRE.     

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。     

不同降水条件下荒漠草原植物的养分含量及回收对增温和氮素添加的响应

为了探讨荒漠草原植物养分回收特征对长期增温和氮素添加的响应以及自然降水变异对其的调控作用,该研究依托实施12年的模拟增温和氮素添加实验平台,在相对多雨的2016年(超过长期均值52%)和相对少雨的2017年(低于长期均值16%),以常见C_3植物银灰旋花(Convolvulus ammannii)和C_4植物木地肤(Kochia prostrata)为研究对象,测定分析绿叶和枯叶的氮磷含量及回收效率。结果表明:(1)在相对多雨年(2016年),增温使2种植物的绿叶氮、枯叶氮、绿叶磷、枯叶磷含量分别增加了14.32%、25.45%、17.97%和46.47%,氮、磷回收效率分别显著减小了9.41%和16.99%(P0.05);氮素添加使2种植物的绿叶氮、枯叶氮、绿叶磷、枯叶磷含量分别提高了17.32%、25.62%、20.21%和51.41%,而氮、磷回收效率显著降低了9.33%和18.89%(P0.05);增温+氮素添加共同处理显著增加了植物氮磷含量、降低了氮磷回收效率。(2)在相对少雨年(2017年),增温、氮素添加、增温+氮素添加处理对植物叶片氮磷含量、回收效率均无显著影响。(3)叶片氮磷含量在物种间差异极显著(P0.000 1),而氮磷回收效率在物种间无显著差异。(4)回归分析表明,植物叶片氮磷含量随着土壤无机氮、有效磷及含水量的增加而增加,植物氮磷回收效率则随着土壤养分和水分的可利用性的增加而降低。研究认为,荒漠草原植物养分回收对全球变化的响应受自然降水变异的调控。     

Influence of frost on nutrient resorption during leaf senescence in a mangrove at its latitudinal limit of distribution

The latitudinal distribution of mangrove species is limited mainly by low temperature. Leaf scorch and massive leaf fall are the predominant symptoms of frost damage. Nutrient resorption during leaf senescence is an important adaptation mechanism of mangroves. Abnormal defoliation disturbs nutrient resorption. We evaluated the effects of frost on nutrient loss of mangroves and the protective effects of warmer seawater inundation on reducing nutrient loss. On January 14, 2009, the most cold-tolerant mangrove Kandelia obovata at its naturally latitudinal limit (Fuding, China, 27°17??N) was exposed to freezing temperature (?2.4°C) for 4 h (minimum ?2.8°C). The freezing air temperature occurred during flood tide, resulting that the flooded shoots were protected by warmer seawater. Frost caused 31.3% and 13.0% defoliation on the exposed shoots and the flooded shoots, respectively. Frost restricted nutrient resorption during leaf senescence. K. obovata resorbed 61% N and 42% P during normal leaf senescence, respectively. However, frost-damaged leaves only resorbed 13% N and 10% P during the course, respectively. Foliar N:P molar ratios were <31, suggesting N limitation. Tidal inundation can partially protect mangroves from frost damage. Reduced nutrient resorption efficiency and massive leaf fall caused by frost add pressure to mangroves under nutrient limitation at their latitudinal limits.     

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.     

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.     

Combined effects of nitrogen addition and litter manipulation on nutrient resorption of Leymus chinensis in a semi‐arid grassland of northern China

Plant growth in semi‐arid ecosystems is usually severely limited by soil nutrient availability. Alleviation of these resource stresses by fertiliser application and aboveground litter input may affect plant internal nutrient cycling in such regions. We conducted a 4‐year field experiment to investigate the effects of nitrogen (N) addition (10 g N·m^?2·year^?1) and plant litter manipulation on nutrient resorption of Leymus chinensis, the dominant native grass in a semi‐arid grassland in northern China. Although N addition had no clear effects on N and phosphorus (P) resorption efficiencies in leaves and culms, N fertilisation generally decreased leaf N resorption proficiency by 54%, culm N resorption proficiency by 65%. Moreover, N fertilisation increased leaf P resorption proficiency by 13%, culm P resorption proficiency by 20%. Under ambient or enriched N conditions, litter addition reduced N and P resorption proficiencies in both leaves and culms. The response of P resorption proficiency to litter manipulation was more sensitive than N resorption proficiency: P resorption proficiency in leaves and culms decreased strongly with increasing litter amount under both ambient and enriched N conditions. In contrast, N resorption proficiency was not significantly affected by litter addition, except for leaf N resorption proficiency under ambient N conditions. Furthermore, although litter addition caused a general decrease of leaf and culm nutrient resorption efficiencies under both ambient and enriched N conditions, litter addition effects on nutrient resorption efficiency were much weaker than the effects of litter addition on nutrient resorption proficiency. Taken together, our results show that leaf and non‐leaf organs of L. chinensis respond consistently to altered soil N availability. Our study confirms the strong effects of N addition on plant nutrient resorption processes and the potential role of aboveground litter, the most important natural fertiliser in terrestrial ecosystems, in influencing plant internal nutrient cycling.     

Modest enhancement of nitrogen conservation via retranslocation in response to gradients in N supply and leaf N status

Plant nutrient resorption, a ubiquitous mechanism of nutrient conservation, has often been proposed to be more pronounced in infertile than fertile habitats, and in species common to infertile compared to fertile habitats, because of the presumed advantage when nutrients are scarce. However, previous studies provide weak and inconsistent empirical support for these hypotheses, although few have examined intraspecific variation across well-quantified resource gradients. This study addresses intraspecific patterns of nutrient resorption for eight species across two N availability gradients on similar soils in an N-limited oak savanna ecosystem: a long-term fire frequency gradient with a negatively correlated N fertility gradient and a long-term N fertilization gradient. We hypothesized that both resorption proficiency (the minimum nutrient level retained in a senesced leaf) and efficiency (the proportional change in leaf nutrient concentration) would decrease with increasing soil N availability and plant N status. For the seven non-N fixers, either resorption proficiency or efficiency decreased modestly in treatments with higher N availability. In contrast, the legume Amorpha canescens Pursh had higher N levels in green and senesced leaves, and resorbed N much more weakly than the non-fixers, and did not respond in terms of proficiency or efficiency to soil N availability. Across all species and sites in each N fertility gradient, a scaling analysis showed greater resorption efficiency in plants with lower N concentrations. Our data suggest that species can have modest resorption responses reflective of soil nutrient availability and differences in resorption related to their N economy that represent mechanisms of nutrient conservation in nutrient-limited soils.     

Nutrient resorption of two evergreen shrubs in response to long-term fertilization in a bog

Plant resorption of multiple nutrients during leaf senescence has been established but stoichiometric changes among N, P and K during resorption and after fertilization are poorly understood. We anticipated that increased N supply would lead to further P limitation or co-limitation with N or K [i.e. P-(co)limitation], decrease N resorption and increase P and K resorption, while P and K addition would decrease P and K resorption and increase N resorption. Furthermore, Ca would accumulate while Mg would be resorbed during leaf senescence, irrespective of fertilization. We investigated the effect of N, P and K addition on resorption in two evergreen shrubs (Chamaedaphne calyculata and Rhododendron groenlandicum) in a long-term fertilization experiment at Mer Bleue bog, Ontario, Canada. In general, N addition caused further P-(co)limitation, increased P and K resorption efficiency but did not affect N resorption. P and K addition did not shift the system to N limitation and affect K resorption, but reduced P resorption proficiency. C. calyculata resorbed both Ca and Mg while R. groenlandicum resorbed neither. C. calyculata showed a higher resorption than R. groenlandicum, suggesting it is better adapted to nutrient deficiency than R. groenlandicum. Resorption during leaf senescence decreased N:P, N:K and K:P ratios. The limited response of N and K and the response of P resorption to fertilization reflect the stoichiometric coupling of nutrient cycling, which varies among the two shrub species; changes in species composition may affect nutrient cycling in bogs.     

Response of leaf, sheath and stem nutrient resorption to 7 years of N addition in freshwater wetland of Northeast China

Background and Aims

Increased N availability induced by agricultural fertilization applications and atmospheric N deposition may affect plant nutrient resorption in temperate wetlands. However, the relationship between nutrient resorption and N availability is still unclear, and most studies have focused on leaf nutrient resorption only. The aim of our study was to examine the response of leaf and non-leaf organ nutrient resorption to N enrichment in a temperate freshwater wetland.

Methods

We conducted a 7-year N addition experiment to investigate the effects of increased N loading on leaf, sheath and stem nutrient (N and P) resorption of two dominant species (Deyeuxia angustifolia and Glyceria spiculosa) in a freshwater marsh in the Sanjiang Plain, Northeast China.

Results

Our results showed that, for both leaf and non-leaf organs (sheath and stem), N addition decreased N resorption proficiency and hence increased litter N concentration. Moreover, the magnitude of N addition effect on N resorption proficiency varied with fertilization rates for D. angustifolia sheaths and stems, and G. spiculosa leaves. However, increased N loading produced inconsistent impacts on N and P resorption efficiencies and P resorption proficiency, and the effects only varied with species and plant organs. In addition, N enrichment increased litter mass and altered litter allocation among leaf, sheath and stem.

Conclusions

Our results highlight that leaf and non-leaf organs respond differentially to N addition regarding N and P resorption efficiencies and P resorption proficiency, and also suggest that N enrichment in temperate freshwater wetlands would alter plant internal nutrient cycles and increase litter quality and quantity, and thus substantially influence ecosystem carbon and nutrient cycles.     

Declining soil fertility does not increase leaf lifespan within species: evidence from the Franz Josef chronosequence, New Zealand

Leaf lifespan varies widely among plant species, from a few weeks to >40 years. This variation is associated with differences in plant form and function, and the distribution of species along resource gradients. Longer leaf lifespans increase the residence time of nutrients and are one mechanism by which plants conserve nutrients; consequently, leaf lifespan should increase within species with declining soil nutrient availability. The Franz Josef chronosequence is a series of post-glacial surfaces along which soil fertility declines strongly with increasing soil age. We used this fertility gradient to test whether leaf lifespans of six common indigenous woody species increased as soil nutrient availability declined. Leaf lifespan varied from 12.4 months in Coprosma foetidissima (Rubiaceae) to 47.1 months in Pseudopanax crassifolius (Araliaceae). These leaf lifespans sample 12% of the full range of leaf lifespans reported globally and occupy a relatively conservative portion of global leaf trait space. Contrary to our expectations, leaf lifespan of two species (Pseudopanax crassifolius and Prumnopitys ferruginea) decreased by 44?61% with increasing soil age and there were no other relationships between soil age and leaf lifespan. Across all species, leaf nutrient residence times increased by 85% for N and 90% for P with declining soil fertility, but this was caused by increased nutrient resorption efficiency rather than by increased leaf longevity. These data demonstrate that plants increase leaf nutrient resorption efficiency rather than leaf lifespan as a within-species response to long-term declines in soil fertility.     

Increased precipitation modulates the influence of nitrogen and litter inputs on the nutrient resorption proficiency rather than efficiency of <Emphasis Type="Italic">Leymus chinensis</Emphasis>

Resorption of nutrients from senescing organs is an important conservation mechanism that is usually influenced by the supply of soil nutrients and plant growth requirements. Therefore, it is likely that increases in nitrogen (N), precipitation, and litter could lead to changes in nutrient resorption because of changes in nutrients in the soil and accelerated plant growth in response to the alleviation of water limitations in arid and semiarid environments. In the current study, we investigated the effects of water, N, and litter addition on the nutrient resorption efficiency and proficiency of N and phosphorus (P) in leaves and stems of Leymus chinensis in Inner Mongolia, China. Our results showed that N addition significantly decreased the N resorption efficiency in leaves under water addition, and increased P resorption efficiency under ambient precipitation conditions. There was no apparent influence of either litter or water addition on N and P resorption efficiencies. However, N and litter addition significantly altered N and P resorption proficiencies, and these effects were modulated by water availability. Furthermore, changes in resorption proficiencies were mainly associated with alterations in the nutritional status of green organs in response to water, N and litter addition, except for leaf P. Our findings highlight the importance of increased precipitation in modulating the nutrient resorption proficiency of plants under potentially increased nutrient availability in semiarid grasslands. Therefore, global changes in precipitation and N, and corresponding litter changes could result in complex effects on plant nutrient economies and, in turn, could influence the return of nutrients to the soil.     

Correction of leaf nutrient resorption efficiency on the mass basis

植物叶片养分重吸收效率的质量损失校正 养分重吸收是植物保持养分的关键机制，但以往多数研究未考虑叶片衰老过程中的质量损失，低估了植物的叶片养分重吸收效率(NuRE)；或只能基于文献中植物功能群水平的平均质量损失，校正所研究的不同物种的叶片NuRE，从而影响了该参数的精确性。本研究通过采集中国北方地区35种常见木本植物的绿色叶片和凋落叶片样本，测量绿色叶片和凋落叶片的质量，计算了叶片衰老过程中的质量损失，并给出了这些物种凋落叶片的质量损失校正系数(MLCF)。总体而言，植物凋落叶片的质量比成熟绿 色叶片平均损失17%，物种水平质量损失变化范围为1.3%–36.8%，功能群水平为11.7%–19.6%。相 应地，这35种木本植物的凋落叶片的MLCF值平均为0.832，变化范围为0.632–0.987。与校正前相 比，用MLCF校正后的NuRE总体上显著增加。例如，校正后氮和磷的平均NuRE皆增高约9%，即比其校正前(低估)更接近真实值；当植物NuRE较低时，这种校正作用(改善)表现得尤为明显。本研究基于物种水平的野外实测数据，报道了中国北方部分常见木本植物的MLCF参考值，研究结果将有助于更准确地计算此类植物的养分重吸收效率，提升本地区植物-土壤系统中养分流通评估的精准度。     

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 conservation increases with latitude of origin in European<Emphasis Type="Italic"> Pinus sylvestris</Emphasis> populations

Nutrient availability varies across climatic gradients, yet intraspecific adaptation across such gradients in plant traits related to internal cycling and nutrient resorption remains poorly understood. We examined nutrient resorption among six Scots pine (Pinus sylvestris L.) populations of wide-ranging origin grown under common-garden conditions in Poland. These results were compared with mass-based needle N and P for 195 Scots pine stands throughout the species' European range. At the common site, green needle N (r(2)=0.81, P=0.01) and P (r(2)=0.58, P=0.08) concentration increased with increasing latitude of population origin. Resorption efficiency (the proportion of the leaf nutrient pool resorbed during senescence) of N and P of Scots pine populations increased with the latitude of seed origin (r(2) > or = 0.67, P < or = 0.05). The greater resorption efficiency of more northerly populations led to lower concentrations of N and P in senescent leaves (higher resorption proficiency) than populations originating from low latitudes. The direction of change in these traits indicates potential adaptation of populations from northern, colder habitats to more efficient internal nutrient cycling. For native Scots pine stands, results showed greater nutrient conservation in situ in cold-adapted northern populations, via extended needle longevity (from 2 to 3 years at 50 degrees N to 7 years at 70 degrees N), and greater resorption efficiency and proficiency, with their greater resorption efficiency and proficiency having genotypic roots demonstrated in the common-garden experiment. However, for native Scots pine stands, green needle N decreased with increasing latitude (r(2)=0.83, P=0.0002), and P was stable other than decreasing above 62 degrees N. Hence, the genotypic tendency towards maintenance of higher nutrient concentrations in green foliage and effective nutrient resorption, demonstrated by northern populations in the common garden, did not entirely compensate for presumed nutrient availability limitations along the in situ latitudinal temperature gradient.     

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.     

Mapping nutrient resorption efficiencies of subarctic cryptogams and seed plants onto the Tree of Life

Nutrient resorption from senescing photosynthetic organs is a powerful mechanism for conserving nitrogen (N) and phosphorus (P) in infertile environments. Evolution has resulted in enhanced differentiation of conducting tissues to facilitate transport of photosynthate to other plant parts, ultimately leading to phloem. Such tissues may also serve to translocate N and P to other plant parts upon their senescence. Therefore, we hypothesize that nutrient resorption efficiency (RE, % of nutrient pool exported) should correspond with the degree of specialization of these conducting tissues across the autotrophic branches of the Tree of Life. To test this hypothesis, we had to compare members of different plant clades and lichens within a climatic region, to minimize confounding effects of climatic drivers on nutrient resorption. Thus, we compared RE among wide‐ranging basal clades from the principally N‐limited subarctic region, employing a novel method to correct for mass loss during senescence. Even with the limited numbers of species available for certain clades in this region, we found some consistent patterns. Mosses, lichens, and lycophytes generally showed low RE_N (<20%), liverworts and conifers intermediate (40%) and monilophytes, eudicots, and monocots high (>70%). RE_P appeared higher in eudicots and liverworts than in mosses. Within mosses, taxa with more efficient conductance also showed higher RE_N. The differences in RE_N among clades broadly matched the degree of specialization of conducting tissues. This novel mapping of a physiological process onto the Tree of Life broadly supports the idea that the evolution of conducting tissues toward specialized phloem has aided land plants to optimize their internal nitrogen recycling. The generality of evolutionary lines in conducting tissues and nutrient resorption efficiency needs to be tested across different floras in different climatic regions with different levels of N versus P availability.     

Is N‐resorption efficiency related to secondary compounds and leaf longevity in coexisting plant species of the arid Patagonian Monte,Argentina?

Leaf longevity and nutrient resorption efficiency are important strategies to conserve plant nutrients. Theory suggests a negative relationship between them and also proposes that high concentration of phenolics in long‐lived leaves may reduce nutrient resorption. In order to provide new evidence on these relationships, we explored whether N‐resorption efficiency is related to leaf longevity, secondary compounds and other leaf traits in coexisting plant species of different life forms in the arid Patagonian Monte, Argentina. We assessed N‐resorption efficiency, green leaf traits (leaf mass per area (LMA), leaf longevity and lignin, total soluble phenolics and N concentrations) and N concentration in senescent leaves of 12 species of different life forms (evergreen shrubs, deciduous shrubs and perennial grasses) with contrasting leaf traits. We found that leaf longevity was positively correlated to LMA and lignin, and negatively correlated to N concentration in green leaves. N concentrations both in green and senescent leaves were positively related. N‐resorption efficiency was not associated with the concentration of secondary compounds (total soluble phenolics and lignin) but it was negatively related to LMA and leaf longevity and positively related to N concentration in green leaves. Furthermore, leaf traits overlapped among life forms highlighting that life forms are not a good indicator of the functional properties (at least in relation to nutrient conservation) of species. In conclusion, our findings indicated that differences in N‐resorption efficiency among coexisting species were more related to N concentration in green leaves, leaf lifespan and LMA than to the presence of secondary compounds at least those assessed in our study (soluble phenolics and lignin). Accordingly, N‐resorption efficiency seems to be modulated, at least in part, by the productivity–persistence trade‐off.     

Drought-deciduous behavior reduces nutrient losses from temperate deciduous trees under severe drought

Nutrient resorption from senescing leaves is an important mechanism of nutrient conservation in temperate deciduous forests. Resorption, however, may be curtailed by climatic events that cause rapid leaf death, such as severe drought, which has been projected to double by the year 2100 in the eastern United States. During a record drought in the southeastern US, we studied 18 common temperate winter-deciduous trees and shrubs to understand how extreme drought affects nutrient resorption of the macronutrients N, P, K, and Ca. Four species exhibited drought-induced leaf senescence and maintained higher leaf water potentials than the remaining 14 species (here called drought-evergreen species). This strategy prevented extensive leaf desiccation during the drought and successfully averted large nutrient losses caused by leaf desiccation. These four drought-deciduous species were also able to resorb N, P, and K from drought-senesced leaves, whereas drought-evergreen species did not resorb any nutrients from leaves lost to desiccation during the drought. For Oxydendrum arboreum, the species most severely affected by the drought, our results indicate that trees lost 50% more N and P due to desiccation than would have been lost from fall senescence alone. For all drought-deciduous species, resorption of N and P in fall-senesced leaves was highly proficient, whereas resorption was incomplete for drought-evergreen species. The lower seasonal nutrient losses of drought-deciduous species may give them a competitive advantage over drought-evergreen species in the years following the drought, thereby impacting species composition in temperate deciduous forests in the future.