Photosynthetic acclimation to light in woody and herbaceous species: a comparison of leaf structure, pigment content and chlorophyll fluorescence characteristics measured in the field

Acclimation of foliage photosynthetic properties occurs with varying time kinetics, but structural, chemical and physiological factors controlling the kinetics of acclimation are poorly understood, especially in field environments. We measured chlorophyll fluorescence characteristics, leaf total carotenoid (Car), chlorophyll (Chl) and nitrogen (N) content and leaf dry mass per area (LMA) along vertical light gradients in natural canopies of the herb species, Inula salicina and Centaurea jacea, and tree species, Populus tremula and Tilia cordata, in the middle of the growing season. Presence of stress was assessed on the basis of night measurements of chlorophyll fluorescence. Our aim was to compare the light acclimation of leaf traits, which respond to light availability at long (LMA and N), medium (Chl a/b ratio, Car/Chl ratio) and short time scales (fluorescence characteristics). We found that light acclimation of nitrogen content per unit leaf area (N(area)), chlorophyll content per unit dry mass (Chl(mass)) and Chl/N ratio were related to modifications in LMA. The maximum PSII quantum yield (F(v) /F(m)) increased with increasing growth irradiance in I. salicina and P. tremula but decreased in T. cordata. Leaf growth irradiance, N content and plant species explained the majority of variability in chlorophyll fluorescence characteristics, up to 90% for steady-state fluorescence yield, while the contribution of leaf total carotenoid content was generally not significant. Chlorophyll fluorescence characteristics did not differ strongly between growth forms, but differed among species within a given growth form. These data highlight that foliage acclimation to light is driven by interactions between traits with varying time kinetics.     

Relationships between leaf nitrogen and limitations of photosynthesis in canopies of Solidago altissima

Vertical distribution patterns of light, leaf nitrogen, and leaf gas exchange through canopies of the clonal perennial Solidago altissima were studied in response to mowing and fertilizer application in a field experiment. Consistent with the distribution of light, average leaf nitrogen content followed a `smooth' exponential decline along the fertilized stands both in control and mown plots. The nitrogen profile along the unfertilized stands in mown plots, however, was `disrupted' by high-nitrogen leaves at the top of shorter ramets that only reached intermediate strata of the canopies. Hence, in these stands leaf nitrogen was significantly increased in short ramets compared with tall ramets for a given light environment, suggesting suboptimal stand structure but not necessarily suboptimal single-ramet architecture. However, at least under the climatic conditions observed during measurements, such disrupture had no substantial effect on stand productivity: model calculations showed that vertical distribution patterns of leaf nitrogen along ramets only marginally influenced the photosynthetic performance of ramets and stands. This is explained by the observed photosynthesis-nitrogen relationship: the rate of photosynthesis per unit amount of leaf nitrogen did not increase with leaf nitrogen content even under saturating light levels indicating that leaf photosynthesis was not nitrogen limited during the measurement periods. Nevertheless, our study indicates that consideration of how architecture(s) of adjacent individual plants interact could be essential for a better understanding of the trade-offs between individual and canopy characteristics for maximizing carbon gain. Such trade-offs may end up in a suboptimal canopy structure, which could not be predicted and understood by classical canopy optimization models.     

Extinction risk of Chinese angiosperms varies between woody and herbaceous species

Aim

Understanding how species' traits and environmental contexts relate to extinction risk is a critical priority for ecology and conservation biology. This study aims to identify and explore factors related to extinction risk between herbaceous and woody angiosperms to facilitate more effective conservation and management strategies and understand the interactions between environmental threats and species' traits.

Location

China.

Taxon

Angiosperms.

Methods

We obtained a large dataset including five traits, six extrinsic variables, and 796,118 occurrence records for 14,888 Chinese angiosperms. We assessed the phylogenetic signal and used phylogenetic generalized least squares regressions to explore relationships between extinction risk, plant traits, and extrinsic variables in woody and herbaceous angiosperms. We also used phylogenetic path analysis to evaluate causal relationships among traits, climate variables, and extinction risk of different growth forms.

Results

The phylogenetic signal of extinction risk differed among woody and herbaceous species. Angiosperm extinction risk was mainly affected by growth form, altitude, mean annual temperature, normalized difference vegetation index, and precipitation change from 1901 to 2020. Woody species' extinction risk was strongly affected by height and precipitation, whereas extinction risk for herbaceous species was mainly affected by mean annual temperature rather than plant traits.

Main conclusions

Woody species were more likely to have higher extinction risks than herbaceous species under climate change and extinction threat levels varied with both plant traits and extrinsic variables. The relationships we uncovered may help identify and protect threatened plant species and the ecosystems that rely on them.     

亚热带常绿林不同冠层小枝叶面积-叶生物量关系研究

叶面积与叶生物量的关系对于理解植物叶片的碳收益和投资权衡策略具有重要意义。收益递减假说认为植物的叶面积与叶生物量成显著异速生长关系,其异速生长指数<1.0,但该假说是否适用于不同生活型(常绿与落叶)亚热带木本植物不同冠层高度(上下冠层)当年生小枝的叶片仍不清楚。以江西亚热带常绿阔叶林的69种常绿与落叶木本植物当年生小枝上的叶为研究对象,采用标准化主轴回归估计(standardized major axis estimation, SMA)方法检验不同冠层高度和生活型叶面积与叶生物量的异速生长关系。结果显示:(1)当年生小枝叶生物量在不同冠层高度和生活型的植物中无显著差异(P>0.05),叶面积在常绿和落叶植物中有显著差异(P<0.05),常绿和落叶植物的比叶重存在显著差异(P<0.05),而落叶植物的比叶重在不同冠层高度存在显著差异(P<0.05),同一冠层,常绿植物比叶重显著高于落叶植物(P<0.05);(2)69种植物的叶面积与叶生物量异速生长指数具有物种特异性,60.9%的物种叶面积与叶生物量呈等速生长关系;(3)不同冠层和生活型植物的叶面积与...     

Relative growth rate of leaf biomass and leaf nitrogen content in several mediterranean woody species

In many plant species, herbivory is a major determinant of leaf mortality and it can cause a strong reduction in productive potential. Most predation occurs on young, expanding leaves. Thus, a rapid growth of the leaves can reduce the impact of predation. Furthermore, in cold Mediterranean climates the length of the growing season is constrained to a short period in spring and early summer owing both to low winter temperatures and drought stress in early summer. Therefore, a rapid deployment of leaf area and a high photosynthetic capacity during the spring and early summer might have important positive effects on the final carbon balance of the leaf population. Relative growth rates (RGR) of leaf biomass were measured in 19 woody species typical of Central Western Spain with deciduous and evergreen habits. Highly significant differences were detected in the leaf growth rate of the different species. The differences between species, however, did not correlate either with the mean leaf life-span of each of the species or with other leaf traits such as photosynthetic capacity, specific leaf area or nitrogen content. Leaf growth rate was positively correlated with time elapsed between leaf initiation and fruit maturation, so that species with fruit dispersal in spring and early summer in general had lower leaf growth rates than species with autumn fruit shedding. This relationship shows the effects of the concurrence between vegetative and reproductive organs for nutrients and other resources. Nitrogen concentration in the leaves was very high at the time of bud break, and declined during leaf expansion owing to the dilution associated with the increase in structural components. The rate of nitrogen dilution was, thus, positively related to the leaf growth rate. Relative growth rates calculated for nitrogen mass in leaves were very low compared to the growth in total mass. This suggests that most leaf nitrogen is translocated from the plant stores to the leaf biomass before the start of leaf expansion and that the contribution of root uptake during leaf expansion is comparatively low.     

Cyanobacterial soil crusts and woody shrub canopies in Kalahari rangelands

Intensive grazing of Kalahari rangelands has led to woody plant encroachment, notably of Acacia mellifera and Grewia flava. The mechanisms causing this process, and the ecological stability of woody plant encroached ecosystems, remain uncertain. Past studies suggest that canopy–soil relations may enhance woody plant competitive dominance. This study aims to investigate one element of this ecological change by examining the spatial distribution of cyanobacterial soil crusts in two vegetation sub‐habitats at sites of different disturbance. Crust burial by litter was also assessed to analyse the dynamics of canopy–crust relations. Our results show there is enhanced cyanobacterial crust cover under A. mellifera canopies and that unlike G. flava canopies, the crust cover remains under A. mellifera even at highly disturbed sites. This canopy–crust association suggests A. mellifera encroachment exhibits intrinsic resilience because of the crusts ability to stabilize the soil surface and increase nutrient retention. Crust burial by litter that accumulates under larger woody plants restricts crust development under canopies. Disturbance restricts crust development in plant interspaces and under G. flava. These two mechanisms combine to restrict crust development to an observed 40% threshold, with nonlinear models required to explain spatial patterns of crust dynamics.     

Critical water contents at leaf,stem and root level leading to irreversible drought-induced damage in two woody and one herbaceous species

Plant water content is a simple and promising parameter for monitoring drought-driven plant mortality risk. However, critical water content thresholds leading to cell damage and plant failure are still unknown. Moreover, it is unclear whether whole-plant or a specific organ water content is the most reliable indicator of mortality risk. We assessed differences in dehydration thresholds in leaf, stem and root samples, hampering the organ-specific rehydration capacity and increasing the mortality risk. We also tested eventual differences between a fast experimental dehydration of uprooted plants, compared to long-term water stress induced by withholding irrigation in potted plants. We investigated three species with different growth forms and leaf habits i.e., Helianthus annuus (herbaceous), Populus nigra (deciduous tree) and Quercus ilex (evergreen tree). Results obtained by the two dehydration treatments largely overlapped, thus validating bench dehydration as a fast but reliable method to assess species-specific critical water content thresholds. Regardless of the organ considered, a relative water content value of 60% induced significant cell membrane damage and loss of rehydration capacity, thus leading to irreversible plant failure and death.     

Acclimation of photosynthetic capacity to irradiance in tree canopies in relation to leaf nitrogen concentration and leaf mass per unit area

The observation of acclimation in leaf photosynthetic capacity to differences in growth irradiance has been widely used as support for a hypothesis that enables a simplification of some soil‐vegetation‐atmosphere transfer (SVAT) photosynthesis models. The acclimation hypothesis requires that relative leaf nitrogen concentration declines with relative irradiance from the top of a canopy to the bottom, in 1 : 1 proportion. In combination with a light transmission model it enables a simple estimate of the vertical profile in leaf nitrogen concentration (which is assumed to determine maximum carboxylation capacity), and in combination with estimates of the fraction of absorbed radiation it also leads to simple ‘big‐leaf’ analytical solutions for canopy photosynthesis. We tested how forests deviate from this condition in five tree canopies, including four broadleaf stands, and one needle‐leaf stand: a mixed‐species tropical rain forest, oak (Quercus petraea (Matt.) Liebl), birch (Betula pendula Roth), beech (Fagus sylvatica L.) and Sitka spruce (Picea sitchensis (Bong.) Carr). Each canopy was studied when fully developed (mid‐to‐late summer for temperate stands). Irradiance (Q, µmol m^?2 s^?1) was measured for 20 d using quantum sensors placed throughout the vertical canopy profile. Measurements were made to obtain parameters from leaves adjacent to the radiation sensors: maximum carboxylation and electron transfer capacity (V_a, J_a, µmol m^?2 s^?1), day respiration (R_da, µmol m^?2 s^?1), leaf nitrogen concentration (N_m, mg g^?1) and leaf mass per unit area (L_a, g m^?2). Relative to upper‐canopy values, V_a declined linearly in 1 : 1 proportion with N_a. Relative V_a also declined linearly with relative Q, but with a significant intercept at zero irradiance (P < 0·01). This intercept was strongly related to L_a of the lowest leaves in each canopy (P < 0·01, r² = 0·98, n= 5). For each canopy, daily lnQ was also linearly related with lnV_a(P < 0·05), and the intercept was correlated with the value for photosynthetic capacity per unit nitrogen (PUN: V_a/N_a, µmol g^?1 s^?1) of the lowest leaves in each canopy (P < 0·05). V_a was linearly related with L_a and N_a(P < 0·01), but the slope of the V_a : N_a relationship varied widely among sites. Hence, whilst there was a unique V_a : N_a ratio in each stand, acclimation in V_a to Q varied predictably with L_a of the lowest leaves in each canopy. The specific leaf area, L_m(cm² g^?1), of the canopy‐bottom foliage was also found to predict carboxylation capacity (expressed on a mass basis; V_m, µmol g^?1 s^?1) at all sites (P < 0·01). These results invalidate the hypothesis of full acclimation to irradiance, but suggest that L_a and L_m of the most light‐limited leaves in a canopy are widely applicable indicators of the distribution of photosynthetic capacity with height in forests.     

Plastid structure and chlorophyll content in leaf primordia of onion bulbs

Angiosperms do not synthesize chlorophyll in the dark. Here we show that leaf primordia in onion bulbs are green, though they developing in dark conditions. We present results that show plastids in green primordia are chloroplasts, and that they contain chlorophyll as well as embryos in seeds of certain angiosperms.     

Thermal acclimation of leaf respiration varies between legume and non-legume herbaceous

Aims Ubiquitous thermal acclimation of leaf respiration could mitigate the respiration increase. However, whether species of different plant functional groups showing distinct or similar acclimation justifies the simple prediction of respiratory carbon (C) loss to a warming climate.     

Evaluating the relationship between leaf chlorophyll concentration and SPAD-502 chlorophyll meter readings

Relationships between chlorophyll concentration ([chl]) and SPAD values were determined for birch, wheat, and potato. For all three species, the relationships were non-linear with an increasing slope with increasing SPAD. The relationships for birch and wheat were strong (r ² ∼ 0.9), while the potato relationship was comparatively weak (r ² ∼ 0.5). Birch and wheat had very similar relationships when the chlorophyll concentration was expressed per unit leaf area, but diverged when it was expressed per unit fresh weight. Furthermore, wheat showed similar SPAD–[chl] relationships for two different cultivars and during two different growing seasons. The curvilinear shape of the SPAD–[chl] relationships agreed well with the simulated effects of non-uniform chlorophyll distribution across the leaf surface and multiple scattering, causing deviations from linearity in the high and low SPAD range, respectively. The effect of non-uniformly distributed chlorophyll is likely to be more important in explaining the non-linearity in the empirical relationships, since the effect of scattering was predicted to be comparatively weak. The simulations were based on the algorithm for the calculation of SPAD-502 output values. We suggest that SPAD calibration curves should generally be parameterised as non-linear equations, and we hope that the relationships between [chl] and SPAD and the simulations of the present study can facilitate the interpretation of chlorophyll meter calibrations in relation to optical properties of leaves in future studies.     

Photosynthetic nitrogen-use efficiency in leaves of woody and herbaceous species

1. Photosynthetic characteristics of an annual herb, Chenopodium album , and an evergreen tree, Quercus myrsinaefolia , were compared to clarify causes of the difference in photosynthetic nitrogen-use efficiency (photosynthetic capacity per unit nitrogen) between leaves of herbaceous and evergreen species.
2. When leaves with the same nitrogen content on an area basis were compared, photosynthetic capacity of C. album was twice as high as that of Q. myrsinaefolia . Gas-exchange measurements showed higher intercellular CO₂ concentration in C. album . Biochemical analyses indicated larger allocation of nitrogen into ribulose-1,5-bisphosphate carboxylase, a key enzyme of photosynthesis, and higher specific activity of ribulose-1,5-bisphosphate carboxylase in C. album . However, these differences were not large.
3. Compositional deviation of ¹³C in leaves of the two species suggested that the drop of CO₂ level between the intercellular space and the chloroplast was slightly larger in Q. myrsinaefolia when compared between the leaves with the same photosynthetic capacity.
4. It is concluded that the difference in photosynthetic nitrogen-use efficiency between C. album and Q. myrsinaefolia is not caused by a sole factor that is markedly different between the two species but by several factors each of which is slightly disadvantageous to Q. myrsinaefolia compared with C. album .     

杭州湾滨海湿地3种草本植物叶片N、P化学计量学的季节变化

动态平衡理论是生态化学计量学的理论基础, 各种有机体是否存在一个固定的化学计量比是生态学研究的热点问题。该文研究了杭州湾滨海湿地3种优势物种海三棱藨草(Scirpus mariqueter)、糙叶薹草(Carex scabrifolia)和芦苇(Phragmites australis)叶片N、P生态化学计量特征的季节变化。结果发现, 3种植物叶片N含量范围分别是7.41-17.12、7.47-13.15和6.03-18.09 mg·g^-1, 平均值(±标准差)分别为(11.69 ± 2.66)、(10.17 ± 1.53)和(11.56 ± 3.19) mg·g^-1; 叶片P范围分别是0.34-2.60、0.41-1.10和0.35-2.04 mg·g^-1, 平均值为(0.93 ± 0.62)、(0.74 ± 0.23)和(0.82 ± 0.53) mg·g^-1; N:P范围分别是7.19-30.63、11.58-16.81和8.62-21.86, 平均值为16.83 ± 8.31、14.53 ± 3.91和16.49 ± 5.51, 可见不同植物其生态化学计量值范围存在一定差异, 但经方差分析发现3种草本植物间生长季节内N、P元素含量差异并不显著(p > 0.05)。各物种叶片N、P含量均表现出在生长初期显著大于其他生长季节(p < 0.05), 生长旺季(6、7月)随着叶片生物量的持续增加, N、P含量逐渐降低并达到最小值, 随后8-9月叶片不再生长而N、P含量逐渐回升, 在10月叶片衰老时N、P含量再次下降; 叶片N:P则在生长初期较小, 在生长旺季先升高后降低, 随后叶片成熟不再生长时又逐渐增加并趋于稳定。     

Facilitation between woody and herbaceous plants that associate with arbuscular mycorrhizal fungi in temperate European forests

In late‐successional environments, low in available nutrient such as the forest understory, herbaceous plant individuals depend strongly on their mycorrhizal associates for survival. We tested whether in temperate European forests arbuscular mycorrhizal (AM) woody plants might facilitate the establishment of AM herbaceous plants in agreement with the mycorrhizal mediation hypothesis. We used a dataset spanning over 400 vegetation plots in the Weser‐Elbe region (northwest Germany). Mycorrhizal status information was obtained from published resources, and Ellenberg indicator values were used to infer environmental data. We carried out tests for both relative richness and relative abundance of herbaceous plants. We found that the subset of herbaceous individuals that associated with AM profited when there was a high cover of AM woody plants. These relationships were retained when we accounted for environmental filtering effects using path analysis. Our findings build on the existing literature highlighting the prominent role of mycorrhiza as a coexistence mechanism in plant communities. From a nature conservation point of view, it may be possible to promote functional diversity in the forest understory through introducing AM woody trees in stands when absent.     

Changes in leaf nitrogen and phosphorus stoichiometry of woody plants along an altitudinal gradient in Changbai Mountain,China

Aims Leaf is the organ of plant photosynthesis, and it is important to understand the drivers for the variations of leaf nitrogen (N) and phosphorus (P) stoichiometry along geographical and climatic gradients. Here we aimed to explore: 1) the changes in leaf nitrogen (N) and phosphorus (P) stoichiometry of woody plants along an altitudinal gradient in Changbai Mountain, and 2) the relative contribution of climate, plant characteristics, and phylogeny to the changes in leaf N, P concentration and N:P.     

Dirunal leaf movement, chlorophyll fluorescence and carbon assimilation in soybean grown under different nitrogen and water availabilities

Diurnal heliotropic leaf movements, photosynthetic gas exchange, and the ratio of variable fluorescence to maximum fluorescence (Fv/Fm) of unrestrained and horizontally restrained leaves from soybean (Glycine max cv. Cumberland) plants grown in two different water and two different nitrogen treatments were measured. Leaves of plants grown in low water or low nitrogen availability treatments displayed more pronounced diaheliotropism (solar tracking) in the afternoon and a longer period of paraheliotropism (light avoiding) at midday relative to those of well-watered, high-nitrogen-grown plants. Photosaturated photosynthetic rates and the photon flux required to saturate photosynthesis were reduced by water stress and nitrogen deficiency. Compared to horizontal leaves, irradiance on orienting leaves was nearer to the breakpoint of the photosynthetic light response curve, where photosynthesis is co-limited by ribulose biphosphate regeneration and carboxylation. This would increase the carbon return on investments of nitrogen into photosynthesis. A positive linear relationship between Fv/Fm and quantum yield of photosynthesis was measured. Leaves of low-nitrogen-grown plants had earlier and more prolonged reductions in Fv/Fm at midday compared to leaves of high nitrogen grown plants of the same water treatment. Within the same water and nitrogen treatment, horizontally restrained leaves had lower midday Fv/Fm in relation to orienting leaves. Nitrogen deficiency and water stress enhanced this difference such that horizontally restrained leaves of low water and low nitrogen grown plants had earlier and longer midday depressions in Fv/Fm.     

Scaling and structure of dicotyledonous leaf venation networks

There have been numerous attempts to derive general models for the structure and function of resource delivery networks in biology. Such theories typically predict the quantitative structure of vascular networks across scales. For example, fractal branching models of plant structure predict that the network dimensions within plant stems or leaves should be scale-free. However, very few empirical examples of such networks are available with which to evaluate such hypotheses. Here, we apply recently developed leaf network extraction software to a global leaf dataset. We find that leaf networks are neither entirely scale-free nor governed entirely by a characteristic scale. Indeed, we find many network properties, such as vein length distributions, which are governed by characteristic scales, and other network properties, notably vein diameter distributions, which are typified by power-law behaviour. Our findings suggest that theories of network structure will remain incomplete until they address the multiple constraints on network architecture.     

A proposal for universal formulas for estimating leaf water status of herbaceous and woody plants based on spectral reflectance properties

The present study deals with the relationships between water status parameters of plant leaves and reflectances (R_λ) at characteristic wavelengths, between 522 and 2450 nm, as well as reflectance ratios, R_λ/R₁₄₃₀, R_λ/R₁₆₅₀, R_λ/R₁₈₅₀, R_λ/R₁₉₂₀, and R_λ/R₁₉₅₀, based on the air-drying experimental results of soybean (Glycine max Merr.), maize (Zea mays L.), tuliptree (Liriodendron tulipifera L.) and viburnum (Viburnum awabuki K. Koch.) plants. The water status parameters include leaf water content per unit leaf area (LWC), specific leaf water content (SWC), leaf moisture percentage of fresh weight (LMP), relative leaf water content (RWC) and relative leaf moisture percentage on fresh weight basis (RMP). Effective spectral reflectances and reflectance ratios for estimating the LWC, SWC, LMP, RWC and RMP were identified. With these spectral indices, approaches to estimating LWC, RWC and RMP were discussed. Eventually, an attempt on universal formulas was made for estimating the leaf moisture conditions of both herbaceous and woody plants as mentioned above. Moreover, applicability of these formulas was checked with the field experimental results of soybean and maize grown under water and nutrient stresses. This revised version was published online in June 2006 with corrections to the Cover Date.