Carbon isotope discrimination and growth response of old Pinus ponderosa trees to stand density reductions

Stand density reductions have been proposed as a method by which old‐growth ponderosa pine (Pinus ponderosa) forests of North America can be converted back to pre‐1900 conditions, thereby reducing the danger of catastrophic forest fires and insect attacks while increasing the productivity of the remaining old‐growth individuals. However, the duration of productivity response of individual trees and the physiological mechanisms underlying such a response remain speculative issues, particularly in old trees. Tree‐ring measurements of carbon isotope ratios (δ¹³C) and basal area increment (BAI) were used to assess the response of intrinsic water‐use efficiency (the ratio of photosynthesis, A to stomatal conductance, g) and growth of individual> 250‐year‐old‐ponderosa pine trees to stand density reductions. It was hypothesized that reductions in stand density would increase soil moisture availability, thus decreasing canopy A/g and increasing carbon isotope discrimination (Δ). Cellulose‐δ¹³C of annual tree rings, soil water availability (estimated from pre‐dawn leaf water potential), photosynthetic capacity, stem basal growth and xylem anatomy were measured in individual trees within three pairs of thinned and un‐thinned stands. The thinned stands were treated 7 to 15 years prior to measurement. The values of δ¹³C and BAI were assessed for 20 consecutive years overlapping the date of thinning in a single intensively studied stand, and was measured for 3 years on either side of the date of thinning for the two other stands to assess the generality of the response. After thinning, Δ increased by 0.89‰ (± 0.15‰). The trees in the un‐thinned stands showed no change in Δ (0.00‰ ± 0.04‰). In the intensively studied trees, significant differences were expressed in the first growing season after the thinning took place but it took 6 years before the full 0.89‰ difference was observed. BAI doubled or tripled after disturbance, depending on the stand, and the increased BAI lasted up to 15 years after thinning. In the intensively studied trees, the BAI response did not begin until 3 years after the Δ response, peaked 1 year after the Δ peak, and then BAI and Δ oscillated in unison. The lag between BAI and Δ was not due to slow changes in anatomical properties of the sapwood, because tracheid dimensions and sapwood‐specific conductivity remained unchanged after disturbance. The Δ response of thinned trees indicated that A/g decreased after thinning. Photosynthetic capacity, as indexed by foliar nitrogen ([N]) and by the relationship between photosynthesis and internal CO₂ (A–C_i curves), was unchanged by thinning, confirming our suspicion that the decline in A/g was due to a relatively greater increase in g in comparison with A. Model estimates agreed with this conclusion, predicting that g increased by nearly 25% after thinning relative to a 15% increase in A. Pre‐dawn leaf water potential averaged 0.11 MPa (± 0.03 MPa) less negative for the thinned compared with the un‐thinned trees in all stands, and was strongly correlated with Δ post‐thinning (R² = 0.91). There was a strong relationship between BAI and modelled A, suggesting that changes in water availability and g have a significant effect on carbon assimilation and growth of these old trees. These results confirm that stand density reductions result in increased growth of individual trees via increased stomatal conductance. Furthermore, they show that a physiological response to stand density reductions can last for up to 15 years in old ponderosa pines if stand leaf area is not fully re‐established.     

Variation in leaf area index and stand leaf mass of European beech across gradients of soil acidity and precipitation

Leaf area index (LAI, the one-sided foliage area per unit ground surface area) is a key determinant of plant productivity which has a large influence on water and energy exchange between vegetation and the atmosphere. The variation in forest LAI across landscapes and environmental gradients and its causes are not sufficiently understood. We measured the LAI of European beech (Fagus sylvatica) by litter trapping in 23 closed, mature stands across gradients of rainfall and soil acidity or fertility. With a mean LAI of 7.4 m² m⁻² (minimum: 5.6, maximum: 9.5 m² m⁻²), beech stands maintained a comparably high leaf area index with relatively small variation along steep environmental gradients. Contrary to expectation, decreasing water availability (rainfall gradient from 1030 to 520 mm yr⁻¹) or increasing soil acidity (pH 3–7) had no significant effect on LAI. Stand leaf mass (M _l) increased slightly with soil fertility (C/N ratio, base saturation). We regressed parameters of site water availability (rainfall), soil fertility or acidity (pH, base saturation, C/N ratio, exchangeable Mg and Al content), and stand structure (stand age and stem density) against LAI and M _l in order to detect environmental controls of stand leaf area. Stand age was the most influential factor for both LAI and M _l (negative relationship). Stem density and the base saturation of the soil affected M _l significantly, but had a weak influence on LAI. We conclude that the leaf area index of beech is mainly under control of age-related physiological factors, whereas the influence of soil chemistry and rainfall is comparably low.     

Forest growth along a rainfall gradient in Hawaii: Acacia koa stand structure,productivity, foliar nutrients,and water- and nutrient-use efficiencies

We tested whether variation in growth of native koa (Acacia koa) forest along a rainfall gradient was attributable to differences in leaf area index (LAI) or to differences in physiological performance per unit of leaf area. Koa stands were studied on western Kauai prior to Hurricane Iniki, and ranged from 500 to 1130 m elevation and from 850 to 1800 mm annual precipitation. Koa stands along the gradient had basal area ranging from 8 to 42 m²/ha, LAI ranging from 1.4 to 5.4, and wood increment ranging from 0.7 to 7.1 tonnes/ha/year. N, P, and K contents by weight of sun leaves (phyllodes) were negatively correlated with specific leaf mass (SLM, g m^-2) across sites; on a leaf area basis, N increased whereas P and K decreased with SLM. LAI, aboveground woody biomass increment, and production per unit leaf area (E) increased as phyllode ¹³C became more negative. The ¹³C data suggested that intrinsic water-use efficiency (ratio of assimilation to conductance) increased as water availability decreased. In five of the six sites, phyllode P contents increased as LAI increased, but biomass increment and E were not correlated with phyllode nutrient contents, suggesting that productivity was limited more by water than by nutrient availability. Because vapor pressure deficits increased with decreasing elevation, actual water-use efficiency (ratio of assimilation to transpiration) was lower at drier, low-elevation sites. There was a trade-off between intrinsic water-use efficiency and production per unit of canopy N or P across the gradient. In summary, koa responds to water limitation both by reducing stand LAI and by adjusting gas exchange, which results in increased intrinsic water-use efficiency but decreased E.     

Leaf Size and Leaf Area Index in <Emphasis Type="Italic">Fagus sylvatica</Emphasis> Forests: Competing Effects of Precipitation,Temperature, and Nitrogen Availability

Plants across diverse biomes tend to produce smaller leaves and a reduced total leaf area when exposed to drought. For mature trees of a single species, however, the leaf area–water supply relationship is not well understood. We tested the paradigm of leaf area reduction upon drought by a transect study with 14 mature Fagus sylvatica forests along a steep precipitation gradient (970–520 mm y⁻¹) by applying two independent methods of leaf size determination. Contrary to expectation, average leaf size in dry stands (520–550 mm y⁻¹) was about 40% larger and SLA was higher than in moist stands (910–970 mm y⁻¹). As a result of increased leaf sizes, leaf area index significantly increased from the high- to the low-precipitation stands. Multiple regression analyses suggested that average leaf size was primarily controlled by temperature, whereas the influence of soil moisture and soil C/N ratio was low. Summer rainfall of the preceding year was the most significant predictor of total leaf number. We assume that leaf expansion of beech was independent of water supply, because it takes place in May with ample soil water reserves along the entire transect. In contrast, bud formation, which determines total leaf number, occurs in mid-summer, when droughts are severest. We conclude that leaf expansion and stand leaf area of beech along this precipitation gradient are not a simple function of water availability, but are controlled by several abiotic factors including spring temperature and possibly also nitrogen supply, which both tend to increase toward drier sites, thus overlaying any negative effect of water shortage on leaf development.     

The 'hydrology' of leaves: co-ordination of structure and function in temperate woody species

The hydraulic conductance of the leaf lamina (K_lamina) substantially constrains whole‐plant water transport, but little is known of its association with leaf structure and function. K_lamina was measured for sun and shade leaves of six woody temperate species growing in moist soil, and tested for correlation with the prevailing leaf irradiance, and with 22 other leaf traits. K_lamina varied from 7.40 × 10^?5 kg m^?2 s^?1 MPa^?1 for Acer saccharum shade leaves to 2.89 × 10^?4 kg m^?2 s^?1 MPa^?1 for Vitis labrusca sun leaves. Tree sun leaves had 15–67% higher K_lamina than shade leaves. K_lamina was co‐ordinated with traits associated with high water flux, including leaf irradiance, petiole hydraulic conductance, guard cell length, and stomatal pore area per lamina area. K_lamina was also co‐ordinated with lamina thickness, water storage capacitance, 1/mesophyll water transfer resistance, and, in five of the six species, with lamina perimeter/area. However, for the six species, K_lamina was independent of inter‐related leaf traits including leaf dry mass per area, density, modulus of elasticity, osmotic potential, and cuticular conductance. K_lamina was thus co‐ordinated with structural and functional traits relating to liquid‐phase water transport and to maximum rates of gas exchange, but independent of other traits relating to drought tolerance and to aspects of carbon economy.     

Water supply regulates structure, productivity, and water use efficiency of Acacia koa forest in Hawaii

We studied changes in stand structure, productivity, canopy development, growth efficiency, and intrinsic water use efficiency (WUE=photosynthesis/stomatal conductance) of the native tree koa (Acacia koa) across a gradient of decreasing rainfall (2600–700 mm) with increasing elevation (700–2000 m) on the island of Hawaii. The stands were located on organic soils on either smooth (pahoehoe) or rough (aa) lava flows. In the greenhouse, we also examined growth and WUE responses to different water regimes of koa seedlings grown from seeds collected in the study area. We tested the hypotheses that (1) stand basal area, aboveground net primary productivity (ANPP), leaf area index (LAI), and growth per unit leaf area decreased with decreasing rainfall, (2) WUE increased with decreasing rainfall or water supply, and (3) WUE responses were caused by stomatal limitation rather than by nutrient limitations to photosynthesis. The carbon isotope composition of phyllode tissues (δ¹³C) was examined as an integrated measure of WUE. Basal area and LAI of koa stands on both pahoehoe and aa lava flows, and ANPP on aa lava flows decreased with elevation. Basal area, LAI, and ANPP of koa in mixed stands with the exotic tropical ash (Fraxinus udhei) were lower compared to single-species koa stands at similar elevations. Along the gradient, phyllode δ¹³C (and therefore WUE) increased with elevation from –30.2 to –26.8‰. Koa in mixed stands exhibited higher (less negative) δ¹³C than in single-species stands suggesting that koa and tropical ash competed for water. In the greenhouse, we observed the same trend observed in the field, as phyllode δ¹³C increased from –27.7 to –24‰ as water supply decreased. Instantaneous gas exchange measurements in the greenhouse showed an inverse correlation of both maximum (morning) photosynthesis (A) and conductance (g) with δ¹³C values and, also, a good agreement between instantaneous (A/g) and integrated measures of WUE. Phyllode δ¹³C was not correlated with foliar concentrations of N or other nutrients in either the field or the greenhouse, indicating that differences in δ¹³C were caused by stomatal limitation rather than by nutrient-related changes in photosynthetic capacity. This study provided evidence that long-term structural and growth adjustments as well as changes in WUE are important mechanisms of koa response to water limitation.     

The effect of fertilization on sap flux and canopy conductance in a Eucalyptus saligna experimental forest

Land devoted to plantation forestry (50 million ha) has been increasing worldwide and the genus Eucalyptus is a popular plantation species (14 million ha) for its rapid growth and ability to grow well on a wide range of sites. Fertilization is a common silvicultural tool to improve tree growth with potential effects on stand water use, but the relationship between wood growth and water use in response to fertilization remains poorly quantified. Our objectives in this study were to determine the extent, timing and longevity of fertilization effects on water use and wood growth in a non‐water limited Eucalyptus saligna experimental forest near Hilo, HI. We evaluated the short‐ and long‐term effects of fertilization on water use by measuring sap flux per unit sapwood area, canopy conductance, transpiration per unit leaf area and water‐use efficiency in control and fertilized stands. Short‐term effects were assessed by comparing sap flux before and after fertilizer application. Long‐term effects were assessed by comparing control plots and plots that had received nutrient additions for 5 years. For the short‐term response, total water use in fertilized plots increased from 265 to 487 mm yr^?1 during the 5 months following fertilization. The increase was driven by an increase in stand leaf area accompanied by an increase in sap flux per unit sapwood area. Sap flux per unit leaf area and canopy conductance did not differ during the 5 months following fertilizer additions. For the last 2 months of our short‐term measurements, fertilized trees used less water per unit carbon gain (361 compared with 751 kg H₂O kg C^?1 in control stands). Trees with 5 years of fertilization also used significantly more water than controls (401 vs. 302 mm yr^?1) because of greater leaf area in the fertilized stands. Sap flux per unit sapwood area, sap flux per unit leaf area, and canopy conductance did not differ between control and fertilized trees in the long‐term plots. In contrast to the short‐term response, the long‐term response of water use per unit wood growth was not significant. Overall, fertilization of E. saligna at our site increased stand water use by increasing leaf area. Fertilized trees grew more wood and used more water, but fertilization did not change wood growth per unit water use.     

热岛效应下亚热带城市植被叶气孔权衡特征及其与叶功能性状的关系

叶片气孔不仅是植物平衡光合-蒸腾关系的重要门户,也是影响大气碳循环与水循环的关键结构。分析热岛效应下福州市乔木、灌木、草本3种生活型和常绿、落叶2种叶习性植物的气孔性状间的差异及其与其他叶功能性状间的权衡关系有助于探究不同类型植物在热环境下的适应策略。以福州市区的自然和半自然植被为研究对象,测定441个植物样本的气孔特征、化学计量特征和形态特征,结果表明：（1）3种生活型、2种叶习性植物的气孔长度（SL）、气孔密度（SD）差异显著（P<0.05）,潜在气孔导度指数（PCI）不存在显著差异（P>0.05）。草本的SL高于灌木和乔木,乔木的SD最高,灌木次之,草本最低;落叶植物的SL高于常绿植物,SD低于常绿植物。（2）SL与SD间的权衡关系稳定存在于3种生活型和2种叶习性植物中,且随着不同生活型和落叶习性植物的生态策略而呈现各异的权衡特征,即当SL一定时,乔木的SD最大,灌木的SD最小,常绿植物的SD大于落叶植物。（3）气孔性状和叶片形态、化学计量特征紧密联系,SL与比叶面积（SLA）正相关（P<0.01）,与叶面积（LA）负相关（P<0.01）;SD与叶氮含量（LNC）、叶磷含量（LPC）、SLA负相关（P<0.01）,与LA正相关（P<0.01）;PCI与LNC、SLA负相关（P<0.01）,与叶厚度（LT）正相关（P<0.05）。（4）复杂的环境是气孔性状变异的重要驱动因素,SL、PCI均与年均温（MAT）负相关（P<0.05）。     

Structural and functional changes in Nothofagus pumilio forests along an altitudinal gradient in Tierra del Fuego,Argentina

Abstract. Structural (density, height, basal area, above‐ground tree biomass, leaf area index) and functional (leaf phenology, growth rate, fine litter fall, leaf decomposition) traits were quantified in four mature forests of Nothofagus pumilio (lenga) along an altitudinal sequence in Tierra del Fuego, Argentina. Three erect forest stands at 220, 440 and 540m and a krummholz stand at 640 m a.s.l. were selected. Along the altitudinal sequence, stem density increased while DBH, height, biomass, leaf‐size and growth period, mean growth rate and decay rate decreased. Dead stems increased and basal area and fine‐litter fall decreased with an increase in elevation among erect forests, but these trends inverted at krummholz. We suggest that krummholz is not only a morphological response to the adverse climate but is also a life form with functional advantages.     

Bundle sheath extensions affect leaf structural and physiological plasticity in response to irradiance

Coordination between structural and physiological traits is key to plants' responses to environmental fluctuations. In heterobaric leaves, bundle sheath extensions (BSEs) increase photosynthetic performance (light‐saturated rates of photosynthesis, A_max) and water transport capacity (leaf hydraulic conductance, K_leaf). However, it is not clear how BSEs affect these and other leaf developmental and physiological parameters in response to environmental conditions. The obscuravenosa (obv) mutation, found in many commercial tomato varieties, leads to absence of BSEs. We examined structural and physiological traits of tomato heterobaric and homobaric (obv) near‐isogenic lines grown at two different irradiance levels. K_leaf, minor vein density, and stomatal pore area index decreased with shading in heterobaric but not in homobaric leaves, which show similarly lower values in both conditions. Homobaric plants, on the other hand, showed increased A_max, leaf intercellular air spaces, and mesophyll surface area exposed to intercellular airspace (S_mes) in comparison with heterobaric plants when both were grown in the shade. BSEs further affected carbon isotope discrimination, a proxy for long‐term water‐use efficiency. BSEs confer plasticity in traits related to leaf structure and function in response to irradiance levels and might act as a hub integrating leaf structure, photosynthetic function, and water supply and demand.     

Canopy structure and leaf nitrogen distribution in a stand of Lysimachia vulgaris L. as influenced by stand density

Summary A hypothesis that a dense stand should develop a less uniform distribution of leaf nitrogen through the canopy than an open stand to increase total canopy photosynthesis was tested with experimentally established stands of Lysimachia vulgaris L. The effect of stand density on spatial variation of photon flux density, leaf nitrogen and specific leaf weight within the canopy was examined. Stand density had little effect on the value of the light extinction coefficient, but strongly affected the distribution of leaf nitrogen per unit area within a canopy. The open stand had more uniform distribution of leaf nitrogen than the dense stand. However, different light climates between stands explained only part of the variation of leaf nitrogen in the canopy. The specific leaf weight in the canopy increased with increasing relative photon flux density and with decreasing nitrogen concentration.     

Variation in bulk‐leaf 13C discrimination,leaf traits and water‐use efficiency–trait relationships along a continental‐scale climate gradient in Australia

Large spatial and temporal gradients in rainfall and temperature occur across Australia. This heterogeneity drives ecological differentiation in vegetation structure and ecophysiology. We examined multiple leaf‐scale traits, including foliar ¹³C isotope discrimination (Δ¹³C), rates of photosynthesis and foliar N concentration and their relationships with multiple climate variables. Fifty‐five species across 27 families were examined across eight sites spanning contrasting biomes. Key questions addressed include: (i) Does Δ¹³C and intrinsic water‐use efficiency (WUE_i) vary with climate at a continental scale? (ii) What are the seasonal and spatial patterns in Δ¹³C/WUE_i across biomes and species? (iii) To what extent does Δ¹³C reflect variation in leaf structural, functional and nutrient traits across climate gradients? and (iv) Does the relative importance of assimilation and stomatal conductance in driving variation in Δ¹³C differ across seasons? We found that MAP, temperature seasonality, isothermality and annual temperature range exerted independent effects on foliar Δ¹³C/WUE_i. Temperature‐related variables exerted larger effects than rainfall‐related variables. The relative importance of photosynthesis and stomatal conductance (g_s) in determining Δ¹³C differed across seasons: Δ¹³C was more strongly regulated by g_s during the dry‐season and by photosynthetic capacity during the wet‐season. Δ¹³C was most strongly correlated, inversely, with leaf mass area ratio among all leaf attributes considered. Leaf N_mass was significantly and positively correlated with MAP during dry‐ and wet‐seasons and with moisture index (MI) during the wet‐season but was not correlated with Δ¹³C. Leaf P_mass showed significant positive relationship with MAP and Δ¹³C only during the dry‐season. For all leaf nutrient‐related traits, the relationships obtained for Δ¹³C with MAP or MI indicated that Δ¹³C at the species level reliably reflects the water status at the site level. Temperature and water availability, not foliar nutrient content, are the principal factors influencing Δ¹³C across Australia.     

Effects of stand age and tree species on canopy transpiration and average stomatal conductance of boreal forests

We quantified the effect of stand age and tree species composition on canopy transpiration (E_C) by analysing transpiration per unit leaf area (E_L) and canopy stomatal conductance (G_S) for boreal trees comprising a five stand wildfire chronosequence. A total of 196 sap flux sensors were used on 90 trees consisting of Betula papyrifera Marsh (paper birch; present in the youngest stand), Populus tremuloides Michx (quaking aspen), Pinus banksiana Lamb. (jack pine), and Picea mariana (Mill.) (black spruce). While fine roots were positively correlated with stand E_C; leaf area index, basal area, and sapwood area were not. Stands less than 70 years old were dominated by Populus tremuloides and Pinus banksiana and stands greater than 70 years old were composed almost entirely of Picea mariana. As Populus tremuloides and Pinus banksiana increased in size and age, they displayed an increasing sapwood to leaf area ratio (A_S : A_L), a constant minimum leaf water potential (Ψ_L), and a constant proportionality between G_S at low vapour pressure deficit (Dj G_Sref) and the sensitivity of G_S to D (–δ). In contrast, A_S : A_L, minimum Ψ_L, and the proportionally between –δ and G_Sref decreased with height and age in Picea mariana. A G_S model that included the effects of D, A_S : A_L, tree height, and for Picea mariana an increasing soil to leaf water potential gradient with stand age, was able to capture the effects of contrasting hydraulic properties of Picea mariana, Populus tremuloides and Pinus banksiana during stand development after wildfire.     

Leaf mining in the Myrtaceae

Abstract 1. Leaf mining insects feed within plant tissue and thus have close physical associations with their host plants. Little is known about the evolution of relationships between leaf miners and their host species. 2. The prevalence of leaf mining was examined in relation to the phylogeny of a dominant family of Australian plants, the Myrtaceae. Sixty‐eight species from the 10 major phylogenetic groups within the family were sampled in south‐eastern Australia. 3. Fifty‐seven per cent of the plant species examined were found to be mined. Several leaf traits were compared between mined and unmined plant species to investigate the potential role of both physical and nutritional characteristics in determining host‐plant susceptibility to these herbivores. 4. The physical leaf traits measured were leaf area, specific leaf area (SLA), lamina thickness, toughness, and amount of fibre (percentage hemicellulose). The nutritional traits were water content, and concentrations of carbon, nitrogen, total phenols and condensed tannins. Principal component analysis showed mined plant species differed from unmined in terms of several leaf traits. 5. In a post hoc regression weighted for phylogeny, leaf miner presence was significantly positively correlated with plant species having thinner laminas and higher phenol concentrations, and was positively correlated with marginal significance to SLA and water content.     

The temporal response to drought in a Mediterranean evergreen tree: comparing a regional precipitation gradient and a throughfall exclusion experiment

Like many midlatitude ecosystems, Mediterranean forests will suffer longer and more intense droughts with the ongoing climate change. The responses to drought in long‐lived trees differ depending on the time scale considered, and short‐term responses are currently better understood than longer term acclimation. We assessed the temporal changes in trees facing a chronic reduction in water availability by comparing leaf‐scale physiological traits, branch‐scale hydraulic traits, and stand‐scale biomass partitioning in the evergreen Quercus ilex across a regional precipitation gradient (long‐term changes) and in a partial throughfall exclusion experiment (TEE, medium term changes). At the leaf scale, gas exchange, mass per unit area and nitrogen concentration showed homeostatic responses to drought as they did not change among the sites of the precipitation gradient or in the experimental treatments of the TEE. A similar homeostatic response was observed for the xylem vulnerability to cavitation at the branch scale. In contrast, the ratio of leaf area over sapwood area (LA/SA) in young branches exhibited a transient response to drought because it decreased in response to the TEE the first 4 years of treatment, but did not change among the sites of the gradient. At the stand scale, leaf area index (LAI) decreased, and the ratios of stem SA to LAI and of fine root area to LAI both increased in trees subjected to throughfall exclusion and from the wettest to the driest site of the gradient. Taken together, these results suggest that acclimation to chronic drought in long‐lived Q. ilex is mediated by changes in hydraulic allometry that shift progressively from low (branch) to high (stand) organizational levels, and act to maintain the leaf water potential within the range of xylem hydraulic function and leaf photosynthetic assimilation.     

Effect of leaf structure and water status on carbon isotope discrimination in field-grown durum wheat

The relationships between leaf and kernel carbon isotope discriminations (Δ) and several leaf structural parameters that are indicators of photosynthetic capacity were studied in durum wheat grown in the field under three water regimens. A set of 144 genotypes were cultivated in two rain-fed trials, and 125 of these were grown under supplementary irrigation before heading. Total chlorophyll and nitrogen (N) contents, the dry mass per unit leaf area (LDM, the reciprocal of specific leaf area) and carbon isotope discrimination (Δ) were measured in penultimate leaves and Δ of mature kernels was also analysed. Both LDM and N per unit area showed significant (P≤ 0.001) negative correlation (r=–0.60 and r=–0.36, respectively) with leaf Δ in the wettest trial. Little or no correlation was found for any structural parameter and leaf Δ in the rain-fed trials. In contrast, in the two rain-fed trials LDM was the parameter with the strongest positive correlation (P≤ 0.001) with kernel Δ (r= 0.47 and 0.30) and grain yield (r= 0.43 and 0.29), whereas no correlation was found in the irrigation trial. These correlations, rather than representing a causal link between the amount of photosynthetic tissue and Δ, were probably indirect associations caused by a parallel effect of water status and phenology on leaf structure, grain Δ and yield. Correlations across trials (i.e. environments) between leaf structure and either Δ and yield were very high, although also spurious. Our results suggest that LDM should be used to cull segregating population differences in leaf Δ based on the internal photosynthetic capacity only in the absence of drought. Selecting for kernel Δ and grain yield on the basis of LDM is worthwhile for rain-fed trials.     

Global response patterns of plant photosynthesis to nitrogen addition: A meta‐analysis

A mechanistic understanding of plant photosynthetic response is needed to reliably predict changes in terrestrial carbon (C) gain under conditions of chronically elevated atmospheric nitrogen (N) deposition. Here, using 2,683 observations from 240 journal articles, we conducted a global meta‐analysis to reveal effects of N addition on 14 photosynthesis‐related traits and affecting moderators. We found that across 320 terrestrial plant species, leaf N was enhanced comparably on mass basis (N_mass, +18.4%) and area basis (N_area, +14.3%), with no changes in specific leaf area or leaf mass per area. Total leaf area (TLA) was increased significantly, as indicated by the increases in total leaf biomass (+46.5%), leaf area per plant (+29.7%), and leaf area index (LAI, +24.4%). To a lesser extent than for TLA, N addition significantly enhanced leaf photosynthetic rate per area (A_area, +12.6%), stomatal conductance (g_s, +7.5%), and transpiration rate (E, +10.5%). The responses of A_area were positively related with that of g_s, with no changes in instantaneous water‐use efficiency and only slight increases in long‐term water‐use efficiency (+2.5%) inferred from ¹³C composition. The responses of traits depended on biological, experimental, and environmental moderators. As experimental duration and N load increased, the responses of LAI and A_area diminished while that of E increased significantly. The observed patterns of increases in both TLA and E indicate that N deposition will increase the amount of water used by plants. Taken together, N deposition will enhance gross photosynthetic C gain of the terrestrial plants while increasing their water loss to the atmosphere, but the effects on C gain might diminish over time and that on plant water use would be amplified if N deposition persists.     

Leaf lifespan as a determinant of leaf structure and function among 23 amazonian tree species

Summary The relationships between resource availability, plant succession, and species' life history traits are often considered key to understanding variation among species and communities. Leaf lifespan is one trait important in this regard. We observed that leaf lifespan varies 30-fold among 23 species from natural and disturbed communities within a 1-km radius in the northern Amazon basin, near San Carlos de Rio Negro, Venezuela. Moreover, leaf lifespan was highly correlated with a number of important leaf structural and functional characterisues. Stomatal conductance to water vapor (g) and both mass and area-based net photosynthesis decreased with increasing leaf lifespan (r²=0.74, 0.91 and 0.75, respectively). Specific leaf area (SLA) also decreased with increasing leaf lifespan (r²=0.78), while leaf toughness increased (r²=0.62). Correlations between leaf lifespan and leaf nitrogen and phosphorus concentrations were moderate on a weight basis and not significant on an area basis. On an absolute basis, changes in SLA, net photosynthesis and leaf chemistry were large as leaf lifespan varied from 1.5 to 12 months, but such changes were small as leaf lifespan increased from 1 to 5 years. Mass-based net photosynthesis (A/mass) was highly correlated with SLA (r²=0.90) and mass-based leaf nitrogen (N/mass) (r²=0.85), but area-based net photosynthesis (A/area) was not well correlated with any index of leaf structure or chemistry including N/area. Overall, these results indicate that species allocate resources towards a high photosynthetic assimilation rate for a brief time, or provide resistant physical structure that results in a lower rate of carbon assimilation over a longer time, but not both.     

喀斯特石漠化地区土壤养分对泡核桃功能性状的影响

为了探究喀斯特石漠化地区植物叶片功能性状及影响因素,以及揭示其对石漠化环境的适应机理,该文以中国南方喀斯特高原峡谷地区的泡核桃(Juglans sigillata)为对象,揭示土壤养分对叶片结构和光合性状的影响效应。结果表明：(1)泡核桃叶功能性状随石漠化等级增加,叶面积减小,比叶面积增大,叶干物质含量和叶组织密度先降后升,蒸腾速率、胞间CO₂浓度、气孔导度和光能利用率先下降后升高,其他性状变化趋势不显著。(2)冗余分析表明土壤养分能够解释37.4%的光合性状变异与53.4%的结构性状变异,其中全磷和溶解性有机碳对光合性状影响最大,而对结构性状影响最显著的是碱解氮和速效磷。(3)比叶面积分别与叶干物质含量极显著负相关,与净光合速率极显著正相关,叶厚度与叶组织密度极显著负相关,蒸腾速率与胞间CO₂浓度、气孔导度极显著正相关,水分利用速率与蒸腾速率、胞间CO₂浓度、气孔导度极显著负相关,光能利用率与净光合速率显著正相关。研究结果表明,泡核桃为适应喀斯特石漠化的特殊生境采取增强生长功能性状,同时提高资源获取能力的开拓型生长策略...     

The vertical distribution of nitrogen and photosynthetic activity at different plant densities in Carex acutiformis

Shoots of the monocotyledonous perennial Carex acutiformis were grown in open (28 shoots m⁻²) and dense stands (280 shoots m⁻²). For fully grown stands the distribution of relative PPFD and leaf nitrogen per unit leaf area over canopy depth was determined. Light response of photosynthesis was measured on leaf segments sampled at various heights in the stands. Relations between parameters of these curves and leaf nitrogen were investigated. Simulations showed that in the open stand daily canopy photosynthesis was not affected by nitrogen redistribution in the canopy. For the dense stand however, a uniform nitrogen distribution would lead to only 73% of the maximum net carbon gain by the stand under optimal nitrogen distribution. The actual canopy photosynthesis was only 7% less than this theoretical maximum; the actual nitrogen distribution of the dense stand clearly tended to the optimal distribution. The vertical pattern of the nitrogen distribution was to a large extent determined by the minimum leaf nitrogen content. The relatively high minimum leaf nitrogen content found for Carex leaves may perhaps be necessary to maintain the physiological function of the basal parts of the leaves.