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.     

Photosynthesis of Conifers in Relation to Annual Growth Cycles and Dry Matter Production

Observations that deciduous and evergreen conifers growing in Britain show similar annual growth increments, despite marked differences in short-term growth rates, led to a comparative study of seasonal photosynthetic capacity in established trees of four coniferous species with contrasting growth habits. The photo-synthetic data were compared with seasonal changes in environmental parameters and chloroplast ultrastructure. The maximum net photosynthetic rates (at 20°C) recorded for Larix leptolepis were higher than those for the evergreen conifers when expressed on a leaf weight basis but not when expressed per unit leaf area. The photosynthetic efficiency of new needles in the evergreen species showed an overall decline from just after needle maturity until just before budbreak in their second season, after which photosynthetic rates recovered temporarily, approaching previous maximum levels. There was no obvious correlation between seasonal photosynthetic efficiency (at 20°C) on the one hand, and daily air, and (30 cm) ground temperatures on the other, and there was no obvious winter suppression of evergreen photosynthetic rates. Evergreen needles showed starch loss and some membrane changes with the onset of winter, but there was no evidence for wintertime chloroplast clumping or membrane disruption.     

Characterization of the short needle phenomenon in red spruce

Red spruce trees (Picea rubens Sarg.) occasionally produce short twigs bearing short needles. The frequency of short needle cohorts is positively correlated with both elevation and defoliation and they are found in greater numbers on trees that regularly experience winter injury. Short needles are smaller, have lower fresh and dry weights, and reduced volumes compared with normal needles. They have reduced cross-sectional areas due to smaller areas of stelar and mesophyll tissue systems. Individual mesophyll cells, however, have the same cross- and longitudinal sectional areas. On a weight or volume basis, both short and normal needles contain similar amounts of chlorophyll and carotenoid pigments. When pigment concentration was calculated on a unit needle or a needle area base, short needles contain more pigment than normal needles. Short needles appear to have a greater photosynthetic efficiency as determined by fluorescence measurements.     

Effect of light and gibberellic acid on photosynthesis during leaf senescence of alstroemeria cut flowers.

The functioning of the photosynthetic apparatus during leaf senescence was investigated in alstroemeria cut flowers by a combination of gas-exchange measurements and analysis of in vivo chlorophyll fluorescence. Chlorophyll loss in leaves of alstroemeria cut flowers is delayed by light and by a treatment of the cut flowers with gibberellic acid (GA₃). The maximal photosynthesis of the leaves was approximately 6 μmol CO₂ m⁻² s⁻¹ at I 350 μmol m⁻² s⁻¹ (PAR) which is relatively low for intact C₃ leaves. Qualitatively the gas-exchange rates followed the decline in chlorophyll content for the various treatments, i.e. light and GA₃-treatment delayed the decline in photosynthetic rates. However, when chlorophyll loss could not yet be observed in the leaves, photosynthetic rates were already strongly decreased. In vivo fluorescence measurements revealed that the decrease in CO₂ uptake is (partly) due to a decreased electron flow through photosystem II. Furthermore, analysis of the fluorescence data showed a high nonphotochemical quenching under all experimental conditions, indicating that the consumption of reducing power in the Calvin cycle is very low. The chlorophyll, remaining after 9 days incubation of leaves with GA₃ in the dark should be considered as a 'cosmetic' pigment without any function in the supply of assimilates to the flowers.     

Rapid and non-invasive detection of plants senescence using a delayed fluorescence technique.

Senescence is a phase of leaf ontogeny marked by declining photosynthetic activity that is paralleled by a decline in chloroplast function. The photosystem II in a plant is considered to be the primary site where delayed fluorescence (DF) is produced. We report here a simple, rapid, and non-invasive technique for detecting plants senescence based on quantitative measurements of DF. In the experimental study, various senescence symptoms induced by age or hormones were examined in the Catharanthus roseus L. G. Don plants. Detecting the DF emissions from leaves with a home-made DF biosensor enables DF parameters of C. roseus to be produced in a short time. Meanwhile, evaluations of leaves senescence were made from measurements of chlorophyll content, ion leakage, and net photosynthesis rate (Pn) based on the consumption of CO2 in the tested plants. The results of our investigation demonstrate that the changes in DF intensity of green plants can truly reflect the changes in photosynthetic capacity and chlorophyll content during age-dependent and hormone-modulated senescence. Moreover, the DF intensity negatively correlates with ion leakage in both types of senescence. With proper calibration, DF may provide an important approach for monitoring senescence process in vivo and quantitatively evaluating senescence extent. Therefore, a DF technique could be potentially useful for less time-consuming and automated screening of the interesting mutants with genetic modifications that change the plant senescence progress.     

European larch phenology in the Alps: can we grasp the role of ecological factors by combining field observations and inverse modelling?

Vegetation phenology is strongly influenced by climatic factors. Climate changes may cause phenological variations, especially in the Alps which are considered to be extremely vulnerable to global warming. The main goal of our study is to analyze European larch (Larix decidua Mill.) phenology in alpine environments and the role of the ecological factors involved, using an integrated approach based on accurate field observations and modelling techniques. We present 2 years of field-collected larch phenological data, obtained following a specifically designed observation protocol. We observed that both spring and autumn larch phenology is strongly influenced by altitude. We propose an approach for the optimization of a spring warming model (SW) and the growing season index model (GSI) consisting of a model inversion technique, based on simulated look-up tables (LUTs), that provides robust parameter estimates. The optimized models showed excellent agreement between modelled and observed data: the SW model predicts the beginning of the growing season (B(GS)) with a mean RMSE of 4 days, while GSI gives a prediction of the growing season length (L(GS)) with a RMSE of 5 days. Moreover, we showed that the original GSI parameters led to consistent errors, while the optimized ones significantly increased model accuracy. Finally, we used GSI to investigate interactions of ecological factors during springtime development and autumn senescence. We found that temperature is the most effective factor during spring recovery while photoperiod plays an important role during autumn senescence: photoperiod shows a contrasting effect with altitude decreasing its influence with increasing altitude.     

Effects of foliar application of nitrogen on the photosynthetic performance and growth of two fescue cultivars under heat stress

The effects of nitrogen fertilization on the growth, photosynthetic pigment contents, gas exchange, and chlorophyll (Chl) fluorescence parameters in two tall fescue cultivars (Festuca arundinacea cv. Barlexas and Crossfire II) were investigated under heat stress at 38/30 °C (day/night) for two weeks. Shoot growth rate of two tall fescue cultivars declined significantly under heat stress, and N supply can improved the growth rates, especially for the Barlexas. Chl content, leaf net photosynthetic rate, stomatal conductance, water use efficiency, and the maximal efficiency of photosystem 2 photochemistry (F_v/F_m) also decreased less under heat stress by N supply, especially in Crossfire II. Moreover, cultivar variations in photosynthetic performance were associated with their different response to heat stress and nitrogen fertilization, which were evidenced by shoot growth rate and photosynthetic pigment contents.     

Seasonal,diurnal and vertical variation in photosynthetic parameters in Phyllostachys humilis bamboo plants

In recent years, temperate bamboo species have been introduced in Europe for multiple uses such as renewable bio-based materials (wood, composites, fibres, biochemicals…) and numerous ecological functions (soil and water conservation, erosion control, phytoremediation…). Despite their interesting potential, little is known on the ecophysiology of these plants in their new habitat. Therefore, we studied gas exchange parameters on a full soil bamboo plantation of Phyllostachys humilis on a test field in Ireland (Europe). We evaluated the seasonal, diurnal and vertical variation of the parameters of two commonly used photosynthetic models, i.e. the light response curve (LRC) model and the model of Farquhar, von Caemmerer and Berry (FvCB). Furthermore, we tested if there were environmental effects on the photosynthetic parameters of these models and if a correlation between photosynthetic parameters and fluorescence parameters was present, fluorescence parameters can be easily and fast determined. Our results show that the gas exchange parameters do not vary diurnally or vertically. Only seasonal variations were found and should, therefore, be taken into account when using the LRC or FvCB model when modelling canopy growth. Therefore, a big-leaf model or a sunlit-shade model can be used for modelling bamboo growth in Western Europe. There is no straightforward relation between environmental variables and the photosynthetic parameters. Although fluorescence parameters showed a correlation with the photosynthetic parameters, application of such correlation may be limited.     

Different strategies for photoprotection during autumn senescence in maple and oak

During autumn senescence, plants must disassemble the photosynthetic apparatus as nutrients are remobilized from the leaves. The goal of this study was to examine changes in relative abundance of photosynthetic proteins and pigments throughout autumn senescence in order to understand the mechanisms of photoprotection used during this process. We sampled leaves from two deciduous tree species [sugar maple (Acer saccharum Marsh.) and swamp white oak (Quercus bicolor Willd.)] throughout autumn during 2010 and 2013. Chlorophyll fluorescence was measured, thylakoids were isolated for western blotting with antibodies to individual proteins and pigment content was assessed. Both species retained high photochemical efficiency until late autumn and showed earlier onset of degradation of photosystem I relative to photosystem II. The species differed in the timing and pattern of degradation of individual photosynthetic proteins and pigments. In maple, there were increases in anthocyanins, more rapid degradation of light‐harvesting proteins and enrichment of xanthophyll cycle pigments in late autumn. In oak, light‐harvesting proteins were retained in higher abundance throughout autumn, PsbS levels increased during early autumn and lutein was enriched in late autumn samples. The results suggest that the species differ in strategies for photoprotection during autumn senescence.     

Response of senescing rice leaves to flooding stress

Flooding stress (FS) induced changes in pigment and protein contents and in photochemical efficiency of thylakoid membranes of chloroplasts were investigated during senescence of primary leaves of rice seedlings. Leaf senescence was accompanied by loss in 2,6-dichlorophenolindophenol (DCPIP) photoreduction, rate of oxygen evolution, quantum yield of photosystem 2 with an increase in MDA accumulation, and non-photochemical quenching (NPQ) of chlorophyll fluorescence. These changes were further aggravated when the leaves during this period experienced FS. The increase in NPQ value under stress may indicate photosynthetic adaptation to FS.     

A reliable methodology for assessing the in vitro photosynthetic competence of two Brazilian savanna species: Hyptis marrubioides and Hancornia speciosa

Evaluation of photosynthetic efficiency is critical for studies on plant responses to environmental conditions as well as for genotype selection; however, there is a lack of reliable and functional protocols for such assessments of plants cultured in vitro. In this study, we aimed to adapt the conventional methodology for measuring gas exchange of plants grown in vitro to analyze the effects of irradiance, flow rate, and air humidity on the photosynthetic rate in cultured plantlets of two ‘Cerrado’ species, namely Hyptis marrubioides and Hancornia speciosa plantlets. Chlorophyll (chl) a fluorescence and chloroplastidic pigment content were also assessed. The highest photosynthetic rates were observed at a photon flux density of 600 μmol m^?2 s^?1, with tube inlet airflow rates between 100 and 300 mL min^?1 and 80 % relative humidity in the inlet air. The electron transport rate curve, by means of chl a fluorescence, was similar to the photosynthetic rate response curve obtained with the infrared gas analyzer. These results demonstrate that both H. marrubioides and H. speciosa seedlings grown in vitro have a functional photosynthetic apparatus and respond to variations in measurement conditions, exhibiting substantial rates of CO₂ assimilation under saturating irradiance conditions. The methodology proposed here can be adapted and applied to other species growing in vitro.     

Root–shoot communication of the seedlings of Japanese larch and a hybrid species grown in different soil-temperature regimes

We studied the effects of soil temperature (7, 15, and 25°C) on the growth and photosynthesis of seedlings of the Japanese larch (Larix kaempferi) and its hybrid larch (L. gmelinii × L. kaempferi) to simulate early stages of regeneration after disturbance. At a soil temperature of 7°C, the root length per unit root biomass, chlorophyll concentration, and photosynthetic nitrogen-use efficiency (PNUE) were markedly lower in the Japanese larch than in the hybrid larch, which may indicate that the hybrid larch is better at acquiring water and nutrients. At ambient temperatures of 17–25°C, the light-saturated photosynthesis rate (P _sat) of both seedlings grown at a soil temperature of 7°C was lower than at 15 or 25°C. By the 16th week, the needle area, root area, and biomass in seedlings of both types were lower at a soil temperature of 7°C than at soil temperatures of 15 or 25°C. At a soil temperature of 25°C, P _sat and nitrogen uptake were lower in both larch species than at 15°C. The growth of the Japanese larch declined sharply from 15 to 25°C; however, the growth of the hybrid larch decreased only slightly from 15 to 25°C. We conclude that an increased soil temperature may retard larch growth in cold regions, especially in the case of the Japanese larch.     

Estimating canopy photosynthetic parameters in maize field based on multi-spectral remote sensing

Aims Determination of canopy photosynthetic parameters is key to accurate simulation of ecosystem function by using remote sensing methods. Currently, remote estimation of vegetation canopy structure characteristics has been widely adopted. However, directly estimating photosynthetic variables (photosynthetic capacity and efficiency) at canopy scale based on field spectrometry combined with CO₂ flux measurements is rare.
Methods In this study, we remotely estimated solar radiation use efficiency (εN, net ecosystem CO₂ exchange/absorbed photosynthetically active radiation (NEE_CO2/APAR); εG, gross primary productivity/absorbed photosynthetically active radiation (GPP/APAR); α, apparent quantum efficiency) and photosynthetic capacity (P_max) based on in situ measurements of spectral reflectance and ecosystem CO₂ fluxes, along with observational data on micrometeorological factors during the entire growing season for a maize canopy in Northeast China.
Important findings Results showed that the seasonal variations in P_max and α exhibited a single peak; whereas the values of εN and εG were higher at the start of vegetative stage and then rapidly decreased with the development of maize until displaying a single peak at the intermediate and late stages of the growing season, coinciding with the occurrence of peak values in P_max. A comparison was made on the predictive performance based on normalized difference vegetation index (NDVI), ratio vegetation index (RVI), wide dynamic range vegetation index (WDRVI), 2-band enhanced vegetation index (EVI2), and chlorophyll index (CI) in estimating four canopy photosynthetic parameters with any combination of two separate wavelengths at the range of 400–1 300 nm, which showed that EVI2 was most closely and linearly related to photosynthetic capacity and efficiency. This study demonstrates that multi-spectral remote sensing information is sensitive to the variations in canopy photosynthetic parameters in maize field and can be used to quantitatively monitor seasonal dynamics of canopy photosynthesis, and to accurately assess crop productivity and ecosystem CO₂ exchange capacity.     

Age-dependent changes in the functions and compositions of photosynthetic complexes in the thylakoid membranes of Arabidopsis thaliana

Photosynthetic complexes in the thylakoid membrane of plant leaves primarily function as energy-harvesting machinery during the growth period. However, leaves undergo developmental and functional transitions along aging and, at the senescence stage, these complexes become major sources for nutrients to be remobilized to other organs such as developing seeds. Here, we investigated age-dependent changes in the functions and compositions of photosynthetic complexes during natural leaf senescence in Arabidopsis thaliana. We found that Chl a/b ratios decreased during the natural leaf senescence along with decrease of the total chlorophyll content. The photosynthetic parameters measured by the chlorophyll fluorescence, photochemical efficiency (F _v/F _m) of photosystem II, non-photochemical quenching, and the electron transfer rate, showed a differential decline in the senescing part of the leaves. The CO₂ assimilation rate and the activity of PSI activity measured from whole senescing leaves remained relatively intact until 28 days of leaf age but declined sharply thereafter. Examination of the behaviors of the individual components in the photosynthetic complex showed that the components on the whole are decreased, but again showed differential decline during leaf senescence. Notably, D1, a PSII reaction center protein, was almost not present but PsaA/B, a PSI reaction center protein is still remained at the senescence stage. Taken together, our results indicate that the compositions and structures of the photosynthetic complexes are differentially utilized at different stages of leaf, but the most dramatic change was observed at the senescence stage, possibly to comply with the physiological states of the senescence process.     

Insight into the photosynthetic apparatus in evergreen and deciduous European oaks during autumn senescence using OJIP fluorescence transient analysis

Climate change is one of the major issues nowadays, and Mediterranean broadleaf species have been suggested to fill possible future gaps created by climate change in Central European forests. To provide a scientific‐based foundation for such practical strategies, it is important to obtain a general idea about differences and similarities in the physiology of Central European and Mediterranean species. In the present study, we evaluated the onset of leaf senescence of a broad spectrum of oak species under the Central European climate in a common garden experiment. Degradation of the photosynthetic apparatus of evergreen (Quercus ilex, Q. suber), semi‐evergreen (Q. × turneri, Q. × hispanica) and deciduous oaks (Q. robur, Q. cerris, Q. frainetto, Q. pubescens) was monitored as chlorophyll content and analysed chlorophyll fluorescence induction transients. In the deciduous species, a significant decline in chlorophyll content was observed during autumn/winter, with Q. pubescens showing the slowest decline. Analysis of fluorescence induction transients revealed a significant decline in quantum efficiency of the primary photochemistry and reaction centre density and later, a decrease in quantum efficiency of end acceptor reduction. Alterations in fluorescence parameters were compared to the decline in chlorophyll content, which occurred much more slowly than expected from the fluorescence data. The evergreen species showed no decline in chlorophyll content, nor different chlorophyll a fluorescence induction behaviour despite temperature falling below 0 °C. The hybrids showed intermediate behaviour between their parental evergreen and deciduous taxa.     

Seasonal,Diurnal and Vertical Variation of Chlorophyll Fluorescence on Phyllostachys humilis in Ireland

In recent years, temperate bamboo species have been introduced in Europe not only as an ornamental plant, but also as a new biomass crop. To measure adaptation stress of bamboo to the climate of Western Europe, chlorophyll fluorescence was measured on a diurnal and seasonal basis in Ballyboughal, Co. Dublin, Ireland. Measurements were attained on the leaves of each node of Phyllostachys humilis. The most frequently used parameter in chlorophyll fluorescence is the photosynthetic efficiency (Fv/Fm). A seasonal dip - as well as a larger variation - of Fv/Fm in spring compared to the rest of the year was observed. Over the year, the upper leaves of the plant perform better than the bottom leaves. These findings were linked to environmental factors such as light intensity, air temperature and precipitation, as increased light intensities, decreasing air temperatures and their interactions, also with precipitation levels have an effect on the photosynthetic efficiency (Fv/Fm) in these plants.     

A novel insight into the regulation of light-independent chlorophyll biosynthesis in <Emphasis Type="Italic">Larix decidua</Emphasis> and <Emphasis Type="Italic">Picea abies</Emphasis> seedlings

Light-independent chlorophyll (Chl) biosynthesis is a prerequisite for the assembly of photosynthetic pigment–protein complexes in the dark. Dark-grown Larix decidua Mill. seedlings synthesize Chl only in the early developmental stages and their Chl level rapidly declines during the subsequent development. Our analysis of the key regulatory steps in Chl biosynthesis revealed that etiolation of initially green dark-grown larch cotyledons is connected with decreasing content of glutamyl-tRNA reductase and reduced 5-aminolevulinic acid synthesizing capacity. The level of the Chl precursor protochlorophyllide also declined in the developing larch cotyledons. Although the genes chlL, chlN and chlB encoding subunits of the light-independent protochlorophyllide oxidoreductase were constitutively expressed in the larch seedlings, the accumulation of the ChlB subunit was developmentally regulated and ChlB content decreased in the fully developed cotyledons. The efficiency of chlB RNA-editing was also reduced in the mature dark-grown larch seedlings. In contrast to larch, dark-grown seedlings of Picea abies (L.) Karst. accumulate Chl throughout their whole development and show a different control of ChlB expression. Analysis of the plastid ultrastructure, photosynthetic proteins by Western blotting and photosynthetic parameters by gas exchange and Chl fluorescence measurements provide additional experimental proofs for differences between dark and light Chl biosynthesis in spruce and larch seedlings.     

Preservation of integrity and activity of Haberlea rhodopensis photosynthetic apparatus during prolonged light deprivation

The effect of prolonged light deprivation on ultrastructure, pigment composition and functions of photosynthetic apparatus of the resurrection plant Haberlea rhodopensis Friv. (Gesneriaceae) was studied. For this purpose, intact plants were kept in darkness for up to 6 months. Haberlea rhodopensis demonstrated extraordinary ability to preserve its photosynthetic machinery intact despite complete absence of light. During the first 4 weeks of light deprivation, we observed preservation of pigment content, chloroplast ultrastructure and a decrease in the rate of CO(2) assimilation. The signs of dark-induced senescence were observed only after the fourth week. This phase was characterized by decrease of pigment content, partial disintegration of chloroplast ultrastructure and by the development of photosystem II down regulation that includes the increases in non-photochemical fluorescence quenching, qN. In comparison with other plants like common bean and Arabidopsis, the processes of dark-induced senescence were very slow and the plants still can recover even after 6 months of light deprivation. We think these findings can open new opportunities for studying not only dark-induced senescence but also to investigate mechanisms determining tolerance to multiple stresses affecting integrity of photosynthetic apparatus.