Association of red coloration with senescence of sugar maple leaves in autumn

We evaluated the association of red coloration with senescence in sugar maple (Acer saccharum Marsh.) leaves by assessing differences in leaf retention strength and the progression of the abscission layer through the vascular bundle of green, yellow, and red leaves of 14 mature open-grown trees in October 2002. Computer image analysis confirmed visual categorization of leaves as predominantly green, yellow or red, and chemical quantification of leaf pigment concentrations verified that leaf color reflected underlying differences in leaf biochemistry. Significantly lower chlorophyll concentrations within red and yellow leaves indicated that senescence was more advanced in leaves from these color categories relative to green leaves. Among leaf types, only red leaves contained high concentrations of anthocyanins. There were significant differences in leaf retention capacity among color categories, with the petioles of green leaves being the most firmly attached to twigs, followed by red and then yellow leaves. Microscopic analysis indicated that yellow leaves had the most advanced extension of the abscission layer through the vasculature, with green and red leaves having significantly less abscission layer progression than yellow. A more limited progression of the abscission layer through vascular bundles may be evidence of delayed leaf senescence that could extend resorption of mobile leaf constituents. Together, results from this study suggest an association between leaf anthocyanin content and functional delays in senescence.     

Mineral partitioning in the phloem during autumn senescence of beech leaves

Summary During the period of leaf senescence in fall, the minerals Mg, Ca, K, P, Cl, S, and Si were compared for occurrence and density in tissue compartments of leaf blade, petiole, and subtending stem of beech (Fagus sylvatica L.). Measurements were made by energy-dispersive X-ray microanalysis. The plant material was collected on 2,9, 16 and 23 October, and showed green, greenyellow, yellow, and red-brown autumn leaf coloration. Mg, K, and P were retrieved from the leaf blade prior to shedding, and deposited mainly in cortex and pith tissues of the stem. S and Ca remained in the leaf, and Si and Cl appeared to accumulate in the leaf prior to shedding. During the four stages of leaf senescence, the phloem compartments of the petiole showed considerable changes in mineral content. In addition, leaf senescence in several cases was accompanied by ion shifting from symplastic to apoplastic compartments and vice versa.     

Divergent strategies of photoprotection in high-mountain plants

Leaves of high-mountain plants were highly resistant to photoinhibitory damage at low temperature. The roles of different photoprotective mechanisms were compared. Mainly, the alpine species Ranunculus glacialis (L.) and Soldanella alpina were investigated because they appeared to apply greatly divergent strategies of adaptation. The ratio of electron transport rates of photosystem II/photosystem I measured in thylakoids from R. glacialis did not indicate a specific acclimation to high irradiance. Low rates of a chloroplast-mediated inactivation of catalase (EC 1.11.1.6) in red light indicated, however, that less reactive oxygen was released by isolated chloroplasts from R. glacialis than by chloroplasts from lowland plants. Leaves of S. alpina and of Homogyne alpina (L.) Cass, but not those of R. glacialis, had a very high capacity for antioxidative protection, relative to lowland plants, as indicated by a much higher tolerance against paraquat-mediated photooxidative damage and a higher -tocopherol content. Accordingly, ascorbate and glutathione were strongly oxidized and already largely destroyed at low paraquat concentrations in leaves of R. glacialis, but were much less affected in leaves of  S. alpina. Non-radiative dissipation of excitation energy was essential for photoprotection of leaves of  S. alpina and depended on the operation of the xanthophyll cycle. Strong non-photochemical quenching of chlorophyll fluorescence occurred also in R. glacialis leaves at high irradiance, but was largely independent of the presence of zeaxanthin or antheraxanthin. For R. glacialis, photorespiration appeared to provide a strong electron sink and a most essential means of photoprotection, even at low temperature. Application of phosphinothricin, which interferes with photorespiration by inhibition of glutamine synthetase, caused a striking reduction of electron transport through photosystem II and induced marked photoinhibition at both ambient and low temperature in leaves of R. glacialis, while  S. alpina was less affected. Received: 18 March 1998 / Accepted: 7 August 1998     

Changes in autumn senescence in northern hemisphere deciduous trees: a meta-analysis of autumn phenology studies

Background and Aims Many individual studies have shown that the timing of leaf senescence in boreal and temperate deciduous forests in the northern hemisphere is influenced by rising temperatures, but there is limited consensus on the magnitude, direction and spatial extent of this relationship.Methods A meta-analysis was conducted of published studies from the peer-reviewed literature that reported autumn senescence dates for deciduous trees in the northern hemisphere, encompassing 64 publications with observations ranging from 1931 to 2010.Key Results Among the meteorological measurements examined, October temperatures were the strongest predictors of date of senescence, followed by cooling degree-days, latitude, photoperiod and, lastly, total monthly precipitation, although the strength of the relationships differed between high- and low-latitude sites. Autumn leaf senescence has been significantly more delayed at low (25° to 49°N) than high (50° to 70°N) latitudes across the northern hemisphere, with senescence across high-latitude sites more sensitive to the effects of photoperiod and low-latitude sites more sensitive to the effects of temperature. Delays in leaf senescence over time were stronger in North America compared with Europe and Asia.Conclusions The results indicate that leaf senescence has been delayed over time and in response to temperature, although low-latitude sites show significantly stronger delays in senescence over time than high-latitude sites. While temperature alone may be a reasonable predictor of the date of leaf senescence when examining a broad suite of sites, it is important to consider that temperature-induced changes in senescence at high-latitude sites are likely to be constrained by the influence of photoperiod. Ecosystem-level differences in the mechanisms that control the timing of leaf senescence may affect both plant community interactions and ecosystem carbon storage as global temperatures increase over the next century.     

Hydraulic analysis of water flow through leaves of sugar maple and red oak

Leaves constitute a substantial fraction of the total resistance to water flow through plants. A key question is how hydraulic resistance within the leaf is distributed among petiole, major veins, minor veins, and the pathways downstream of the veins. We partitioned the leaf hydraulic resistance (R(leaf)) for sugar maple (Acer saccharum) and red oak (Quercus rubra) by measuring the resistance to water flow through leaves before and after cutting specific vein orders. Simulations using an electronic circuit analog with resistors arranged in a hierarchical reticulate network justified the partitioning of total R(leaf) into component additive resistances. On average 64% and 74% of the R(leaf) was situated within the leaf xylem for sugar maple and red oak, respectively. Substantial resistance-32% and 49%- was in the minor venation, 18% and 21% in the major venation, and 14% and 4% in the petiole. The large number of parallel paths (i.e. a large transfer surface) for water leaving the minor veins through the bundle sheath and out of the leaf resulted in the pathways outside the venation comprising only 36% and 26% of R(leaf). Changing leaf temperature during measurement of R(leaf) for intact leaves resulted in a temperature response beyond that expected from changes in viscosity. The extra response was not found for leaves with veins cut, indicating that water crosses cell membranes after it leaves the xylem. The large proportion of resistance in the venation can explain why stomata respond to leaf xylem damage and cavitation. The hydraulic importance of the leaf vein system suggests that the diversity of vein system architectures observed in angiosperms may reflect variation in whole-leaf hydraulic capacity.     

Age-specific mating strategies and reproductive senescence

While males gain obvious direct advantages from multiple mating, the reproductive capacity of females is more constrained. The reason why polyandry evolved in females is therefore open to many conjectures. One hypothesis postulates that females gain indirect benefits by increasing the probability of siring young from high quality males. To explore this hypothesis, we used the natural variation of the reproductive value that males and females undergo through age. The age-related variation of phenotypic performance might then induce variations in mating strategies in males and females. Using the common lizard (Lacerta vivipara) as our model system, we showed that reproductive immaturity and senescence created variability in both male and female reproductive success (including survival of offspring). Consistent with theory, males at their best-performing phenotype adopted a polygynous strategy. These males were of an intermediate age and they produced offspring of higher viability than younger and older males. In contrast, females at their best performing phenotype, also of an intermediate age, were less polyandrous than other less-performing females. Middle-aged females tended to mate with males of an intermediate age and produced litters with higher viability independently from their reproductive strategy. Males of an intermediate age enhanced their fitness by additional matings with young or old females. Young and old females increased their fitness by being more polyandrous. Polyandry therefore appears as means to seek for good males. A positive correlation between males and their partners' fitness disagree with the idea that polyandry is the result of a sexual conflict in this species.     

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.     

Different roles of alpha- and beta-branch xanthophylls in photosystem assembly and photoprotection

Xanthophylls (oxygenated carotenoids) are essential components of the plant photosynthetic apparatus, where they act in photosystem assembly, light harvesting, and photoprotection. Nevertheless, the specific function of individual xanthophyll species awaits complete elucidation. In this work, we analyze the photosynthetic phenotypes of two newly isolated Arabidopsis mutants in carotenoid biosynthesis containing exclusively alpha-branch (chy1chy2lut5) or beta-branch (chy1chy2lut2) xanthophylls. Both mutants show complete lack of qE, the rapidly reversible component of nonphotochemical quenching, and high levels of photoinhibition and lipid peroxidation under photooxidative stress. Both mutants are much more photosensitive than npq1lut2, which contains high levels of viola- and neoxanthin and a higher stoichiometry of light-harvesting proteins with respect to photosystem II core complexes, suggesting that the content in light-harvesting complexes plays an important role in photoprotection. In addition, chy1chy2lut5, which has lutein as the only xanthophyll, shows unprecedented photosensitivity even in low light conditions, reduced electron transport rate, enhanced photobleaching of isolated LHCII complexes, and a selective loss of CP26 with respect to chy1chy2lut2, highlighting a specific role of beta-branch xanthophylls in photoprotection and in qE mechanism. The stronger photosystem II photoinhibition of both mutants correlates with the higher rate of singlet oxygen production from thylakoids and isolated light-harvesting complexes, whereas carotenoid composition of photosystem II core complex was not influential. In depth analysis of the mutant phenotypes suggests that alpha-branch (lutein) and beta-branch (zeaxanthin, violaxanthin, and neoxanthin) xanthophylls have distinct and complementary roles in antenna protein assembly and in the mechanisms of photoprotection.     

Larger diurnal temperature range undermined later autumn leaf senescence with warming in Europe

Aim

Climate change regulates autumn leaf senescence date (LSD), exhibiting a strong phenological control of plant carbon uptake. Unlike the delaying effect of daily mean temperature (T_mean) on LSD, the impact of warming asymmetry in daytime and nighttime, as evidenced by variations of the diurnal temperature range (DTR), remains elusive. The objectives of this study were to investigate physiological and ecological impacts of DTR on LSD using long-term in situ observations and to predict the future trends of LSD under warming.

Location

Europe.

Time period

1950–2015.

Major taxa studied

Plant phenology.

Methods

We used partial correlation analysis, multiple linear regression and ridge regression to explore the impacts of DTR on LSD. To quantify the importance of potential drivers of LSD, we trained random forest models and applied the SHapley Additive exPlanations method to isolate the marginal contributions of each predictor on LSD. For LSD modelling and projection, we first evaluated two temperature-driven LSD models [i.e., cooling-degree-day (CDD, without DTR effect) and day–night-temperature CDD (DN_CDD, with DTR effect)], then applied them to predict future LSDs.

Results

We found that observational increases in T_mean and DTR had contrasting effects on LSD. Increased T_mean delayed the LSD, whereas larger DTR overall had an advancing effect. Considering the DTR effect, the T_mean sensitivity of LSD was 14% lower than presently estimated (2.4 vs. 2.8 days °C⁻¹). Warming asymmetry-related drought stress and plant functional traits (i.e., plant isohydricity and water-use efficiency) potentially explained the advancing effect of DTR on LSD. We found that current projections of future LSD are overestimated because the DTR effect is discounted, suggesting the need for an adequate understanding of how plant phenology responds to warming asymmetry.

Main conclusions

Our findings highlight the importance of DTR in controlling LSD variations with an advancing-dominant effect and call for the improvement of phenology modelling incorporating the DTR effect. Given that DTR showed a globally narrowing trend over the last several decades, more efforts are needed to understand the potential ecological impacts of warming asymmetry and vegetation response to climate change.     

Pigment dynamics and autumn leaf senescence in a New England deciduous forest,eastern USA

The leaves of woody plants at Harvard Forest in Central Massachusetts, USA, changed color during senescence; 70% (62/89) of the woody species examined anatomically contained anthocyanins during senescence. Anthocyanins were not present in summer green leaves, and appeared primarily in the vacuoles of palisade parenchyma cells. Yellow coloration was a result of the unmasking of xanthophyll pigments in senescing chloroplasts. In nine red-senescing species, anthocyanins were not detectable in mature leaves, and were synthesized de novo in senescence, with less than 20µg cm^–2 of chlorophyll remaining. Xanthophyll concentrations declined in relation to chlorophyll to the same extent in both yellow- and red-leaved taxa. Declines in the maximum photosystemII quantum yield of leaves collected prior to dawn were only slightly less in the red-senescing species, indicating no long-term protective activity. Red-leaved species had significantly greater mass/area and lower chlorophylla/b ratios during senescence. Nitrogen tissue concentrations in mature and senescent leaves negatively correlated to anthocyanin concentrations in senescent leaves, weak evidence for more efficient nitrogen resorption in anthocyanic species. Shading retarded both chlorophyll loss and anthocyanin production in Cornus alternifolia, Acer rubrum, Acer saccharum, Quercus rubra and Viburnum alnifolium. It promoted chlorophyll loss in yellow-senescing Fagus grandifolia. A reduced red:far-red ratio did not affect this process. Anthocyanins did not increase leaf temperatures in Q.rubra and Vaccinium corymbosum on cold and sunny days. The timing of leaf-fall was remarkably constant from year to year, and the order of senescence of individual species was consistent.     

Distinct male reproductive strategies in two closely related oak species

Reproductive strategies of closely related species distributed along successional gradients should differ as a consequence of the trade‐off between competition and colonization abilities. We compared male reproductive strategies of Quercus robur and Q. petraea, two partly interfertile European oak species with different successional status. In the studied even‐aged stand, trees of the late‐successional species (Q. petraea) grew faster and suffered less from intertree competition than trees of the early‐successional species (Q. robur). A large‐scale paternity study and a spatially explicit individual‐based mating model were used to estimate parameters of pollen production and dispersal as well as sexual barriers between species. Male fecundity was found to be dependent both on a tree's circumference and on its environment, particularly so for Q. petraea. Pollen dispersal was greater and more isotropic in Q. robur than in Q. petraea. Premating barriers to hybridization were strong in both species, but more so in Q. petraea than in Q. robur. Hence, predictions based on the competition–colonization trade‐off are well supported, whereas the sexual barriers themselves seem to be shaped by colonization dynamics.     

Differential effects of sugar maple, red oak, and hemlock tannins on carbon and nitrogen cycling in temperate forest soils

Tannins are abundant secondary chemicals in leaf litter that are hypothesized to slow the rate of soil-N cycling by binding protein into recalcitrant polyphenol–protein complexes (PPCs). We studied the effects of tannins purified from sugar maple, red oak, and eastern hemlock leaf litter on microbial activity and N cycling in soils from northern hardwood–conifer forests of the northeastern US. To create ecologically relevant conditions, we applied tannins to soil at a concentration (up to 2 mg g⁻¹ soil) typical of mineral soil horizons. Sugar maple tannins increased microbial respiration significantly more than red oak or hemlock tannins. The addition of sugar maple tannins also decreased gross N mineralization by 130% and, depending upon the rate of application, decreased net rates of N mineralization by 50–290%. At low concentrations, the decrease in mineralization appeared to be driven by greater microbial-N immobilization, while at higher concentrations the decrease in mineralization was consistent with the formation of recalcitrant PPCs. Low concentrations of red oak and hemlock tannins stimulated microbial respiration only slightly, and did not significantly affect fluxes of inorganic N in the soil. When applied to soils containing elevated levels of protein, red oak and hemlock tannins decreased N mineralization without affecting rates of microbial respiration, suggesting that PPC formation decreased substrate availability for microbial immobilization. Our results indicate that tannins from all three species form recalcitrant PPCs, but that the degree of PPC formation and its attendant effect on soil-N cycling depends on tannin concentration and the pool size of available protein in the soil.     

Nitrogen metabolism and remobilization during senescence

Senescence is a highly organized and well-regulated process. As much as 75% of total cellular nitrogen may be located in mesophyll chloroplasts of C(3)-plants. Proteolysis of chloroplast proteins begins in an early phase of senescence and the liberated amino acids can be exported to growing parts of the plant (e.g. maturing fruits). Rubisco and other stromal enzymes can be degraded in isolated chloroplasts, implying the involvement of plastidial peptide hydrolases. Whether or not ATP is required and if stromal proteins are modified (e.g. by reactive oxygen species) prior to their degradation are questions still under debate. Several proteins, in particular cysteine proteases, have been demonstrated to be specifically expressed during senescence. Their contribution to the general degradation of chloroplast proteins is unclear. The accumulation in intact cells of peptide fragments and inhibitor studies suggest that multiple degradation pathways may exist for stromal proteins and that vacuolar endopeptidases might also be involved under certain conditions. The breakdown of chlorophyll-binding proteins associated with the thylakoid membrane is less well investigated. The degradation of these proteins requires the simultaneous catabolism of chlorophylls. The breakdown of chlorophylls has been elucidated during the last decade. Interestingly, nitrogen present in chlorophyll is not exported from senescencing leaves, but remains within the cells in the form of linear tetrapyrrolic catabolites that accumulate in the vacuole. The degradation pathways for chlorophylls and chloroplast proteins are partially interconnected.     

Leaf age and season influence the relationships between leaf nitrogen, leaf mass per area and photosynthesis in maple and oak trees

Abstract. Seasonal changes in photosynthesis, leaf nitrogen (N) contents and leaf mass per area (LMA) were observed over three growing seasons in open-grown sun-lit leaves of red maple ( Acer rubrum ), sugar maple ( A. sacchamm ) and northern pin oak ( Quereus ellipsoidalis ) trees in southern Wisconsin. Net photosynthesis and leaf N were highly linearly correlated on both mass and area bases within all species from late spring until leaf senescence in fall. Very early in the growing season leaves had high N concentrations, but low photosynthetic rates per unit leaf N, suggesting that leaves were not fully functionally developed at that time. Leaf N per unit area and LMA had nonparallel seasonal patterns, resulting in differing relationships between leaf N/area and LMA in the "early versus late growing season. As a result of differences in seasonal patterns between leaf N/area and LMA, net photosynthesis/area was higher for a given LMA in the spring than fall, and the overall relationships between these two parameters were poor.     

伊犁河谷春秋草场草地生态调查及其恢复对策

针对新疆伊犁河谷春秋草场严重退化的现状,通过对退化草场、围栏封育并实施灌溉的草场、单纯围栏的草场和灌溉但不封牧草场样方的对比调查,从地表植物的生物量、植物种类、优质牧草所占比例和草地植物多样性指数等的变化规律和特征方面,探讨了伊犁河谷地区春秋草场退化的原因和恢复措施。结果表明:适当灌溉后草地生物量为1540.5g.m-2,远高于不灌溉的草地(生物量为188.13g.m-2),表明干旱是本区草地植被生长一个重要制约因素;而仅改变草地的水分条件,在超载率达123%的大环境下,草场中优质牧草的比例由封育的62.38%下降到未封育的2.08%。因此,适度放牧并辅以灌溉措施是本区天然草场恢复的2个必要条件。     

Changes in antioxidative enzymes in cucumber cotyledons during natural senescence: comparison with those during dark-induced senescence

Changes in 7 antioxidative enzymes in naturally senescent cotyledons of cucumber ( Cucumis sativus ) were investigated. The activities of superoxide dismutase (SOD; EC 1.15.1.1), catalase (EC 1.11.1.6), dehydroascorbate reductase (EC 1.8.5.1) and glutathione reductase (GR; EC 1.6.4.2) gradually decreased during the progression of senescence, while those of ascorbate peroxidase (APX; EC 1.11.1.11) and guaiacol peroxidase (GPX; EC 1.11.1.7) gradually increased. The activity of monodehydroascorbate reductase (MDAR; EC 1.6.5.4) was not significantly changed. Western blot analysis showed that the protein level of mitochondrial SOD gradually declined. The protein level of catalase transiently decreased and then increased in the later stages of senescence, despite the decrease in its activity. The overall behavior was markedly different from that found in cotyledons of artificially senescing seedlings transferred into darkness; the activities of SOD, catalase, APX, GPX and GR gradually increased.