Interpreting the decrease in leaf photosynthesis during flowering in mango

Little is known about the effect of flowering on leaf photosynthesis. To understand why net photosynthesis (A(net)) is lower in Mangifera indica L. leaves close to inflorescences than in leaves on vegetative shoots, we measured nitrogen and carbohydrate concentrations, chlorophyll a fluorescence and gas exchange in recently matured leaves on vegetative terminals and on floral terminals of 4-year-old trees. We used models to estimate photosynthetic electron fluxes and mesophyll conductance (g(m)). Lower A(net) in leaves close to developing inflorescences was attributable to substantial decreases in stomatal conductance and g(m), and also in photosynthetic capacity as indicated by the decrease in the light-saturated rate of photosynthetic electron transport (J(max)). The decrease in J(max) was the result of decreases in the amount of foliar nitrogen per unit leaf area, and may have been triggered by a decrease in sink activity as indicated by the increase in the hexose:sucrose ratio. Parameters measured on leaves close to panicles bearing set fruits were generally intermediate between those measured on leaves on vegetative shoots and on leaves close to inflorescences, suggesting that the changes in A(net) associated with flowering are reversible.     

Physiological significance of anthocyanins during autumnal leaf senescence

The light screen hypothesis states that foliar anthocyanins shade the photosynthetic apparatus from excess light. In this paper we extend the light screen hypothesis, postulating that plant species at risk of photoinhibitory conditions during autumnal leaf senescence often utilize anthocyanins to protect the photosynthetic apparatus during the period of nutrient resorption. When senescence-related photosynthetic instabilities are compounded by other environmental stresses, particularly low temperature, severe photoinhibition may result in reduced resorption of critical foliar nutrients, which can significantly affect plant fitness. There is evidence that environments where low and often freezing temperatures are common in autumn selectively favor the production of anthocyanins in senescing foliage. The stimuli for, and the timing and location of, autumnal anthocyanin production are all consistent with a photoprotective role for these pigments in senescing leaves. Furthermore, differences in nitrogen allocation strategies between early and late successional species appear to affect photosynthetic stability during leaf senescence, resulting in a reduced need for foliar autumnal anthocyanins in many early successional plants. The ecological and physiological evidence presented in this paper suggest that, for many deciduous species, the production of anthocyanins provides effective photoprotection during the critical period of foliar nutrient resorption.     

Seasonal Changes in the Nutrient Concentrations of Leaves and Leaf Litter in a Young Cryptomeria japonica Stand

The leaves and leaf litter of Cryptomeria japonica D. Don was collected from April 1994 to March 1995 to describe the seasonal changes in nutrient concentrations in leaves and leaf litter. Nitrogen (N), phosphorus (P) and potassium (K) concentrations were in the order new leaves > old leaves > leaf litter, whereas calcium (Ca) concentration was in the order leaf litter > old leaves > new leaves during the whole year. N, P and K concentrations were at their highest during the new leaf growth phase, and then decreased as a result of the diluting effect and translocation, whereas Ca increased with time. Magnesium did not show any clear seasonal trend compared with other nutrients. N resorption efficiency was lower than P resorption efficiency. There were two nutrient resorption peaks, which could be attributed to high nutrient translocation to new leaves in the spring and to translocation from old leaves before senescence in the autumn. A significant correlation between N and P resorption was observed.     

Inhibitory effect of flowering and early fruit growth on leaf photosynthesis in mango

Carbohydrate and nitrogen contents, chlorophyll fluorescence and gas exchange were measured in leaves from both vegetative and reproductive terminal shoots of 12-year-old flowering mango trees. Reproductive shoot leaves were close to swelling floral buds, inflorescences or panicles bearing set fruits. Leaves close to inflorescences had lower rates of mitochondrial respiration (Rd) and net photosynthesis (Anet), and lower stomatal conductance (gs) and quantum efficiency of photosystem II under actinic light than vegetative shoot leaves. Leaf nitrogen concentration, which decreased from the beginning until the end of flowering, was lower in leaves close to inflorescences than in vegetative shoot leaves. However, these differences and changes were counterbalanced by an increase in leaf mass-to-area ratio so that leaf nitrogen per unit leaf area (Na) remained nearly constant during the whole flowering period, except in leaves close to panicles bearing set fruits. Net CO2 assimilation rate simulated by a biochemical model of leaf photosynthesis (Urban et al. 2003) was much higher than Anet measured at an ambient CO2 partial pressure (Ca) of either 36 or 70 Pa. The overestimation of Anet was more pronounced in leaves close to inflorescences, to panicles bearing set fruits and to reversing inflorescences (characterized by the appearance of leaves in terminal positions on inflorescences) than in vegetative shoot leaves. It is concluded that low Anet in leaves close to inflorescences was probably due neither to changes in Na nor to a decrease in Rubisco activity induced by low gs, but rather to a decrease in electron flow in photosystem II. This decrease was not directly associated with higher starch or soluble sugar contents.     

Species differences in timing of leaf fall and foliage chemistry modify nutrient resorption efficiency in deciduous temperate forest stands

Extensive variation in fractional resorption of mineral elements from plant leaves is still not fully understood. In multi-species forest stands, species leaf fall phenology and leaf constitution may significantly modify the timing of nutrient return to the soil and overall plant nutrient loss. We studied leaf fall and nutrient loss kinetics, and leaf composition in three natural, temperate, deciduous broadleaf forest stands to determine the role of timing of leaf abscission and nutrient immobilization in cell walls on nutrient resorption efficiency of senescing leaves. Nitrogen (N), phosphorus and potassium contents decreased continuously in attached leaves after peak physiological activity during mid-season. Changes in nutrient contents of attached leaves were paralleled by decreases in nutrient contents in freshly fallen leaf litter. In different species and for different nutrients, resorption of nutrients from senescing leaves proceeded with different kinetics. The maximum nutrient resorption efficiency (the fraction of specific nutrient resorbed from the leaves at the end of leaf fall) did not depend on the mid-seasonal nutrient concentration. Species with earlier leaf fall resorbed leaf nutrients at a faster rate, partly compensating for the earlier leaf fall. Nevertheless, the litter-mass weighted mean nutrient contents in leaf litter were still larger in species with earlier leaf fall, demonstrating an inherent trade-off between early leaf fall and efficient nutrient resorption. This trade-off was most important for N. Losses of the non-mobile nutrients calcium and magnesium were unaffected by the timing of leaf fall. There was large variation in the maximum N resorption efficiency among species. Correlations among leaf chemical variables suggested that the maximum N resorption efficiency decreased with the increasing fraction of cell walls in the leaves, possibly due to a greater fraction of N occluded in cell wall matrix. We conclude that species leaf fall phenology and leaf chemistry modify the timing and quantities of plant nutrient losses, and that more diverse forest stands supporting a spectrum of species with different phenologies and leaf types produce litter with more variable chemical characteristics than monotypic stands.     

Changes in respiration and chemical content during autumnal senescence of Populus tremuloides and Quercus rubra leaves

Changes in respiration rate, chemical content and chemical concentration were measured in leaves of field-grown Populus tremuloides Michx. and Quercus rubra L. trees throughout the growing season and autumnal senescence. Chlorophyll, soluble sugar, N, P, K and Mg contents and concentrations all declined during leaf senescence, whereas Ca content and concentration increased. Leaf dry mass per area declined 24 and 35% in P. tremuloides and Q. rubra, respectively, during senescence. In leaves of both species, respiration rates peaked during leaf expansion in the spring and then declined, as a result of reduced cytochrome-mediated respiration, to reach relatively constant rates by midsummer. In senescing P. tremuloides leaves, respiration rates remained relatively constant until mid-October and then declined rapidly. In senescing Q. rubra leaves, respiration rates increased in late September, as a result of the appearance of residual respiration that could not be reduced by respiratory inhibitors, and then declined quickly in early November. No changes in alternative pathway respiratory activity were observed in leaves of either species during senescence until late autumn when rates declined. Because respiration rates were correlated with both leaf sugar and nitrogen content during leaf senescence, we conclude that respiration rates were maintained or increased during leaf senescence to supply energy for degradation and mobilization of chemical constituents.     

Transfer of salt and nutrients in Bruguiera gymnorrhiza leaves during development and senescence

The life span of leaves in Bruguiera gymnorrhiza, non-secretor of salt, can be divided into a leaf developing stage, a leaf functioning stage and a leaf senescing stage. The concentrations (mg/g) and the contents (mg/leaf) of Na and Cl increased at the leaf developing stage and remained almost constant at the leaf functioning stage. At the leaf senescing stage, the concentrations of Na and Cl increased markedly by 45 and 31% respectively, while their contents only increased by 16 and 4% respectively. The K/Na ratio remained constant at the leaf functioning stage, and decreased at the leaf senescing stage. During leaf senescence, there was a marked decline in leaf mass (20%) and in leaf area (15%). During senescence, 60% of its N, 48% of its P and 46% of its K was transferred out of the senescing leaf.     

Water relations, gas exchange and growth of dominant and suppressed shoots of Arbutus unedo L

Basal shoots produced by Arbutus unedo L. after cutting at ground level vary in size and growth rate, and are classified accordingly as dominant or suppressed. The suppressed shoots eventually cease growth and die. In this study, we investigated the role of light and water in the competition among shoots of A. unedo. Dominant and suppressed shoots of A. unedo showed similar leaf water potentials and tissue water relations over the year, suggesting that water status is not responsible for the lack of flushing in suppressed shoots. Although suppressed shoots did not flush under low light, they showed many characteristics of shade-tolerant plants. Leaves of suppressed shoots had lower leaf conductance and light-saturated photosynthetic rate, and higher specific leaf area than leaves of dominant shoots. We conclude that light was the main resource determining competition among shoots and the death of suppressed shoots.     

不同林龄樟子松叶片养分含量及其再吸收效率

树木叶片的养分再吸收效率能够反映树木对养分保存、利用以及对养分贫瘠环境的适应能力。以科尔沁沙地东南缘章古台地区樟子松人工林为研究对象,分析了11、20、29、45年生树木叶片的基本特征、养分含量及其再吸收效率。结果表明:叶片衰老后其质量和面积明显减少;叶片凋落前的平均养分含量没有表现出随樟子松年龄增加而出现有规律的变化;凋落叶片中的N、P、K、Mg含量表现出随年龄增加而增加的趋势,而Ca的趋势与之相反;11年生和20年生的樟子松叶片N、P、K的再吸收效率相似,都显著高于29年生和45年生樟子松(P<0.05),而樟子松叶片对Mg的再吸收效率表现出随年龄增大而显著降低,Ca随叶片的衰老而不断累积,再吸收效率表现为负值,20年生的樟子松叶片Ca再吸收效率最大,11年生和45年生最低。樟子松叶片的N、P、K、Mg养分再吸收效率随年龄增加而降低的趋势表明,随年龄增加樟子松对贫瘠养分生境的适应能力逐渐降低,反映了樟子松养分保存方面的衰退特征。     

Gas exchange and leaf aging in an evergreen oak: causes and consequences for leaf carbon balance and canopy respiration

Leaves of Mediterranean evergreens experience large variations in gas exchange rates over their life span due to aging and seasonally changing environmental conditions. Accounting for the changing respiratory physiology of leaves over time will help improve estimations of leaf and whole-plant carbon balances. Here we examined seasonal variations in light-saturated net CO(2) assimilation (A(max)), dark respiration (R(d)) and the proportional change in R(d) per 10 °C change in temperature (Q(10) of R(d)) in previous-year (PY) and current-year (CY) leaves of the broadleaved evergreen tree Quercus ilex L. A(max) and R(d) were lower in PY than in CY leaves. Differences in nitrogen between cohorts only partly explained such differences, and rates of A(max) and R(d) expressed per unit of leaf nitrogen were still significantly different between cohorts. The decline in A(max) in PY leaves did not result in the depletion of total non-structural carbohydrates, whose concentration was in fact higher in PY than CY leaves. Leaf-level carbon balance modeled from gas exchange data was positive at all ages. Q(10) of R(d) did not differ significantly between leaf cohorts; however, failure to account for distinct R(d) between cohorts misestimated canopy leaf respiration by 13% across dates when scaling up leaf measurements to the canopy. In conclusion, the decline in A(max) in old leaves that are close to or exceed their mean life span does not limit the availability of carbohydrates, which are probably needed to sustain new growth, as well as R(d) and nutrient resorption during senescence. Accounting for leaf age as a source of variation of R(d) improves the estimation of foliar respiratory carbon release at the stand scale.     

Girdling effects on fruit set and quantum yield efficiency of PSII in two Citrus cultivars

Girdling effects on fruitlet abscission, leaf chlorophyll, chlorophyll a fluorescence and carbohydrate concentration in various flowering and vegetative shoots were studied during natural fruit drop in two Citrus cultivars. Irrespective of shoot type, girdling delayed fruitlet abscission, but only fruitlets borne on leafy shoots had increased final fruit set. Chlorophyll a fluorescence analysis revealed differences in quantum yield efficiency of photosystem II of light adapted leaves (Phi(PSII)) among shoot types and in response to girdling. In young leaves of vegetative shoots, girdling decreased Phi(PSII), whereas Phi(PSII) increased from Day 30 after girdling in young leaves of leafy flowering shoots; however, Phi(PSII) did not change in mature leaves during fruit set in either control or girdled trees. Girdling altered leaf carbohydrate concentrations and the photosynthetic performance of the various shoot types. Our results indicate that, in Citrus, several carbohydrate-based regulatory mechanisms of photosynthesis coexist during carbohydrate accumulation brought about by girdling. It is concluded that the delay in fruitlet abscission and the increase in Phi(PSII )observed in girdled leafy flowering shoots are the mechanisms underlying the enhancement of fruit set after girdling.     

毛竹叶片衰老过程碳氮元素及灰分含量的季节动态

于2006年4月—2007年3月,对福建永春毛竹(Phyllostachys pubescens)成熟叶和老叶的C、N元素及灰分含量的季节动态及元素的内吸收率进行研究。结果表明:①毛竹成熟叶和老叶的C、N元素含量存在一定的季节变化,从成熟叶到老叶,C含量保持相对稳定,而N含量呈现下降趋势。②毛竹成熟叶、老叶灰分含量月变化趋势相似,毛竹成熟叶、老叶的灰分含量在秋季(10月)至翌年春季(3月)灰分含量较高,而春夏季相对较低;毛竹成熟叶的灰分含量在7.56%～15.91%之间,老叶的灰分含量在7.46%～16.67%,老叶灰分含量高于成熟叶(P=0.0199)。③毛竹叶片衰老过程N的内吸收率为36.68%。     

Leaf senescence and late-season net photosynthesis of sun and shade leaves of overstory sweetgum (Liquidambar styraciflua) grown in elevated and ambient carbon dioxide concentrations

We examined the effects of elevated CO2 concentration ([CO2]) on leaf demography, late-season photosynthesis and leaf N resorption of overstory sweetgum (Liquidambar styraciflua L.) trees in the Duke Forest Free Air CO2 Enrichment (FACE) experiment. Sun and shade leaves were subdivided into early leaves (formed in the overwintering bud) and late leaves (formed during the growing season). Overall, we found that leaf-level net photosynthetic rates were enhanced by atmospheric CO2 enrichment throughout the season until early November; however, sun leaves showed a greater response to atmospheric CO2 enrichment than shade leaves. Elevated [CO2] did not affect leaf longevity, emergence date or abscission date of sun leaves or shade leaves. Leaf number and leaf area per shoot were unaffected by CO2 treatment. A simple shoot photosynthesis model indicated that elevated [CO2] stimulated photosynthesis by 60% in sun shoots, but by only 3% in shade shoots. Whole-shoot photosynthetic rate was more than 12 times greater in sun shoots than in shade shoots. In senescent leaves, elevated [CO2] did not affect residual leaf nitrogen, and nitrogen resorption was largely unaffected by atmospheric CO2 enrichment, except for a small decrease in shade leaves. Overall, elevated [CO2] had little effect on the number of leaves per shoot at any time during the season and, therefore, did not change seasonal carbon gain by extending or shortening the growing season. Stimulation of carbon gain by atmospheric CO2 enrichment in sweetgum trees growing in the Duke Forest FACE experiment was the result of a strong stimulation of photosynthesis throughout the growing season.     

檫木叶片秋季衰老时叶色、色素和营养元素的关系

【目的】从叶片衰老角度研究檫木叶片叶色、色素和营养元素的变化规律,为秋色叶观赏树种檫木的选育以及栽培提供理论依据。【方法】以栽植于同一环境条件下的3年生檫木为试验材料,从檫木叶片停止生长到脱落,分5个时期对叶片的叶色值、色素含量和营养元素含量进行观测和分析。【结果】观测前期与观测后期,檫木叶片叶色值、色素含量和营养元素含量存在极显著差异;叶片进入衰老阶段后,叶绿素含量和类胡萝卜素含量呈下降趋势,而花色素苷含量逐渐上升;N、P、K含量在叶片衰老阶段逐渐下降。由典型相关分析可知,叶色a*值与叶绿素含量具有显著负相关,与花色素苷含量呈显著正相关;花色素苷含量与N含量成反比,类胡萝卜素含量与P含量成正比;N元素再利用效率的载荷值符号与N含量载荷值的符号相反,而P元素再利用效率的载荷值符号与P含量载荷值的符号相同;檫木叶片衰老分为3个阶段,第1阶段从9月中下旬到10月上旬,为叶片衰老准备期;第2阶段从10月中下旬到11月上旬,为叶片缓慢衰老期;第3阶段从11月中下旬到叶片脱落,为叶片衰老末期。【结论】檫木叶色最佳观赏期是从10月中下旬到11月上旬的叶片缓慢衰老期;从叶片衰老准备期到叶片衰老末期,叶绿素和类胡萝卜素被分解,花色素苷合成;N、P、K 3种营养元素逐渐被转移,其中N含量越高,N元素再利用效率就越低;与之相反,P含量越高,P元素再利用效率就越高。     

Determination of lignin,holocellulose, and organic solvent extractives in fresh leaf,litterfall, and organic material on forest floor using near-infrared reflectance spectroscopy

We report the results of a study regarding the near-infrared reflectance spectra of various leaf stages from fresh to senescing, and to decomposing leaf. A broad absorbance feature increased in the region of 1100–1400nm with the advance of the leaf senescence and leaf decomposition. A decrease was seen in the region over 1440nm during the senescence and decomposition process. These differences of spectra showed the changes in constituents of leaf in terms of the degree of the senescence and decomposition. A comparison of multiple linear regression between the near-infrared reflectance spectra and proximate chemical analyses showed that near-infrared reflectance spectroscopy achieved a certain level of useful accuracy. We consider that near-infrared reflectance spectroscopy has the potential to predict the contents of carbon fractions in plant materials, and that this method can replace previous methods due to faster determination of carbon fractions, and its ability to significantly increase the number of samples that can be collected and measured.     

Growth and carbohydrate status of coppice shoots of hybrid poplar following shoot pruning

Fifteen, 1-year-old Populus maximowiczii Henry x P. nigra L. 'MN9' trees were decapitated and allowed to sprout. After 8 weeks, all had 6 to 10 coppice shoots. All shoots, except the tallest (dominant) shoot, were removed from five of the trees (pruned treatment), and shoot growth, gas exchange and carbohydrate status were compared in the pruned and unpruned trees. Although photosynthetic rate of recently mature leaves of pruned trees was approximately 50% greater than that of leaves on the dominant shoot of unpruned trees, and the dry weight of leaves of pruned trees was 37% greater than that of the leaves on the dominant shoot of unpruned trees, the shoot dry matter relative growth rate did not differ between treatments. Concentrations of water-soluble carbohydrates and starch in the uppper stem and leaves of the dominant shoot were similar in pruned and unpruned trees. However, relative to that of the dominant shoot in unpruned trees, the lower stem in pruned trees was depleted in both soluble carbohydrates and starch. Starch deposition, assessed as the quantity of (14)C-starch in tissues 24 h after a fully expanded source leaf was labeled with (14)CO(2), was 3.9 times greater in roots of pruned trees than in roots of unpruned trees. We conclude that early removal of all but the dominant shoot reduces the carbohydrate status of the roots and the lower portion of the stem by eliminating the excised shoots as a source of photosynthate.     

The dynamics of rooting in Triplochiton scleroxylon cuttings: their relation to leaf area, node position, dry weight accumulation, leaf water potential and carbohydrate composition

Single-node, leafy stem cuttings of Triplochiton scleroxylon K. Schum. were collected from successive nodes down the uppermost shoot of 2-shoot stockplants. The leaves were trimmed to 10, 50 and 100 cm(2) before the cuttings were set under intermittent mist to root. Batches of cuttings were harvested after 0, 14, 28 and 42 days to assess leaf water potential, dry weight and carbohydrate content of their leaf and stem portions. Cuttings with leaf areas of 10, 50 and 100 cm(2) increased in total dry weight by 29, 61 and 90%, respectively, during the 6-week period. The increase in dry weight was accompanied by increases in reflux-extracted soluble carbohydrates (RSC), water-soluble carbohydrates (WSC) and starch. By contrast, increase in leaf area reduced leaf water potential of cuttings before root emergence. Fewer large-leaved cuttings rooted than smaller-leaved cuttings, suggesting that rooting ability is at least partially determined by the balance between photosynthesis and transpiration. Fewer roots per cutting were produced on cuttings with 10 cm(2) leaves than on cuttings with larger leaves. Node position affected increments in dry weight, carbohydrate content and leaf water potential, with differences between nodes on day 0 generally being lost or slightly reversed by day 14. Rooting ability was not related to initial (day 0) carbohydrate content, suggesting that rooting is dependent on carbohydrates formed after severance. During the rooting period, the proportions of total non-structural carbohydrate as WSC and starch were reversed, from mostly WSC on day 0 to mostly starch by day 42. These changes in WSC and starch occurred most rapidly in large-leaved cuttings.     

Evaluation of leaf display of evergreen broadleaved tree species and deciduous tree species in warm temperate conifer plantations

We investigated the sapling leaf display in the shade among trees of various leaf lifespans co-occurring under the canopy of a warm-temperate conifer plantation. We measured leaf-area ratio (aLAR) and morphological traits of saplings of evergreen broadleaved tree species and a deciduous tree species. Although we found large interspecific and intraspecific differences in aLAR even among saplings of similar size in the homogeneous light environment, we did not find a consistent trend in aLAR with leaf lifespan among the species. While deciduous trees annually produced a large leaf area, some evergreen broadleaved trees retained their leaves across years and had aLAR values as high as those of deciduous trees. Among leaf-level, shoot-level, and individual-level morphological traits, aLAR was positively correlated with current-year shoots mass per aboveground biomass in deciduous trees, and with the area of old leaves per aboveground mass in evergreen broadleaved trees. Thus, tree-to-tree variation in the degrees of annual shoot production and the accumulation of old leaves were responsible for the interspecific and intraspecific variations in aLAR.     

Partitioning of photoassimilates in avocado (Persea americana Mill.) during flowering and fruit set

To assess competition for photoassimilates among developing inflorescences, fruitlets and vegetative shoots in avocado (Persea americana Mill.), the distribution of recent photoassimilates was examined before and during flowering, fruit set, and the transition from sink to source of developing vegetative shoots. Mature leaves, located proximal to developing reproductive organs, and developing leaves, located distal to those organs, were exposed to a one-hour pulse of (14)CO(2). Translocation of radiolabeled assimilates was monitored over time and among organs. Overall distribution of assimilates was dependent on dry mass of tissues regardless of organ type. Flowers and fruitlets did not demonstrate greater sink strength than non-autotrophic leaves. Organs receiving assimilates were in phyllotactic alignment with source leaves. Photoassimilates were never limiting, although flower and fruitlet abscission occurred during and long after this competitive period. Carbohydrate availability was sufficient to support the growth of both developing fruitlets and leaves during early stages of reproductive development, and it did not limit fruitlet growth or stimulate fruitlet abscission.