Autumn Senescence Research Articles

Dormancy regulator Prunus mume DAM6 promotes ethylene-mediated leaf senescence and abscission.

Leaf senescence and abscission in autumn are critical phenological events in deciduous woody perennials. After leaf fall, dormant buds remain on deciduous woody perennials, which then enter a winter dormancy phase. Thus, leaf fall is widely believed to be linked to the onset of dormancy. In Rosaceae fruit trees, DORMANCY-ASSOCIATED MADS-box (DAM) transcription factors control bud dormancy. However, apart from their regulatory effects on bud dormancy, the biological functions of DAMs have not been thoroughly characterized. In this study, we revealed a novel DAM function influencing leaf senescence and abscission in autumn. In Prunus mume, PmDAM6 expression was gradually up-regulated in leaves during autumn toward leaf fall. Our comparative transcriptome analysis using two RNA-seq datasets for the leaves of transgenic plants overexpressing PmDAM6 and peach (Prunus persica) DAM6 (PpeDAM6) indicated Prunus DAM6 may up-regulate the expression of genes involved in ethylene biosynthesis and signaling as well as leaf abscission. Significant increases in 1-aminocyclopropane-1-carboxylate accumulation and ethylene emission in DEX-treated 35S:PmDAM6-GR leaves reflect the inductive effect of PmDAM6 on ethylene biosynthesis. Additionally, ethephon treatments promoted autumn leaf senescence and abscission in apple and P. mume, mirroring the changes due to PmDAM6 overexpression. Collectively, these findings suggest that PmDAM6 may induce ethylene emission from leaves, thereby promoting leaf senescence and abscission. This study clarified the effects of Prunus DAM6 on autumn leaf fall, which is associated with bud dormancy onset. Accordingly, in Rosaceae, DAMs may play multiple important roles affecting whole plant growth during the tree dormancy induction phase.

No slowdown of growing season extension with warming in a permafrost‐affected meadow on the Tibetan Plateau

Abstract The Tibetan Plateau holds the world's largest alpine permafrost and is undergoing an acceleration of warming. Phenological shifts over alpine permafrost in a warmer world have been little studied and are greatly underrepresented in current syntheses. Here, we conducted seasonal and gradient temperature‐controlled experiments in a permafrost‐affected meadow to evaluate how warming drives shifts in spring and autumn phenology, and associated growing‐season length at both community and species levels. We found that there is no sign of slowdown in spring advance with warming under a higher year‐around warming treatment, aligning with a future medium warming scenario. Although spring advance led to an advance in autumn senescence according to spring‐only warming experiments, the advance could not offset the delay due to concurrent warming. As a result, year‐around warming significantly delayed autumn senescence, although there was a deceleration in delay with warming under high temperature treatment than under the low one. This finding can be attributed to the possibility that winter warming is insufficient to reduce chilling accumulation, which would not delay spring phenology and then lead to a non‐slowdown in spring phenological advancement. Taken together, there is no slowdown in an extension of growing season length with warming under a higher year‐around warming treatment, with an increase of length by 9 and 21 days at the end of this century under a CO2 stabilization and medium warming scenarios, respectively. Synthesis. Our results suggest that a continued growing season extension at least under the medium warming scenario would help permafrost‐affected meadow ecosystems to mitigate permafrost carbon release on the Tibetan Plateau.

Flavonols do not affect aphid load in green or senescing birch leaves but coincide with a decrease in Photosystem II functionality.

Instead of red anthocyanins, birches synthesise colourless (to human eye), UV-absorbing flavonols during autumn senescence. To test if flavonols protect against insects, and if leaves with high or low amounts of flavonols differ in their photosynthetic functions, aphid-free and aphid-infested green and senescing birch leaves were collected from outdoor-grown trees and analysed. Photosynthetic parameters were greatly affected by the leaf chlorophyll content (i.e. the phase of senescence). Photochemical quenching and the amount of functional Photosystem I decreased linearly with chlorophyll content, while FV/FM (Photosystem II functionality) decreased strongly only at the end of senescence. Non-photochemical quenching of excitation energy (NPQ) increased towards the end of senescence. However, no significant differences in the total flavonol amounts, nor in individual flavonol species, were found between aphid-free and aphid-infested leaves, suggesting that flavonols play no role in defence against aphid herbivory. Interestingly, both green and senescing leaves with a high flavonol content showed low FV/FM values. High flavonol content slowed down PSII photoinhibition and improved recovery, but only in green leaves. Previously, we proposed that anthocyanins provide an additional sink for photosynthates at the nitrogen resorption phase during autumn senescence, and the present data may suggest that flavonol synthesis plays a similar role.

Weakened connection between spring leaf-out and autumn senescence in the Northern Hemisphere.

Vegetation autumn phenology is critical in regulating the ecosystem carbon cycle and regional climate. However, the dominant drivers of autumn senescence and their temporal shifts under climate change remain poorly understood. Here, we conducted a multi-factor analysis considering both direct climatic controls and biological carryover effects from start-of-season (SOS) and seasonal peak vegetation activities on the end-of-season (EOS) to fill these knowledge gaps. Combining satellite and ground observations across the northern hemisphere, we found that carryover effects from early-to-peak vegetation activities exerted greater influence on EOS than the direct climatic controls on nearly half of the vegetated land. Unexpectedly, the carryover effects from SOS on EOS have significantly weakened over recent decades, accompanied by strengthened climatic controls. Such results indicate the weakened constraint of leaf longevity on senescence due to prolonged growing season in response to climate change. These findings underscore the important role of biological carryover effects in regulating vegetation autumn senescence under climate change, which should be incorporated into the formulation and enhancement of phenology modules utilized in land surface models.

Multitemporal airborne imaging spectrometry and fluorometry reveal contrasting photoprotective responses of trees

The Photochemical Reflectance Index (PRI) and solar induced fluorescence (SIF) provide information on plant photosynthetic activity. PRI and SIF are both strongly influenced by irradiance, but uncertainties related to the interpretation of these light responses at large spatial scales remain, partly due to a shortage of suitable data from aircraft or satellite platforms. The goal of this study was to explore interpretations of the PRI- and SIF-light responses of trees owing to species, functional types (evergreen and deciduous) and season. Using airborne hyperspectral and ultraspectral imagery in a North American urban forest, we derived PRI, SIF, and albedo (an indicator of illumination) at the 1-m pixel level. We then quantified crown-level PRI and SIF light responses of ten different tree species at three time points from late-summer to autumnal senescence using hierarchical models. Our results confirmed that both PRI and SIF were strongly influenced by illumination with PRI decreasing and SIF increasing with illumination. Both slope and intercept of the PRI-albedo relationship changed with season, but the pattern varied among species and functional types. SIF values decreased during autumnal senescence for all species, but evergreen species exhibited less seasonal decline in the slope of SIF-albedo relationship compared to deciduous species. The PRI and SIF light responses derived from the airborne imagery offer complementary information on dynamic photosynthesis responses presumably due to varying canopy structure, pigmentation and photoprotection among species and functional types. From airborne platforms, PRI- and SIF-light responses can be used to explore the contrasting physiological responses of individual tree crowns, providing a spatially and temporally explicit view of dynamic plant traits related to photoregulation and a novel view of functional diversity for entire landscapes.

Do red and yellow autumn leaves make use of different photoprotective strategies during autumn senescence?

Our goal was to determine whether anthocyanin-producing species (red) use different photoprotective strategies to cope with excess light during fall senescence compared with non-anthocyanin-producing species (yellow). In a previous study, we found that a yellow species retained the photoprotective PsbS protein in late autumn, while a red species did not. Specifically, we tested the hypothesis that red species make less use of zeaxanthin and PsbS-mediated thermal dissipation, as they rely on anthocyanins for photoprotection. We monitored four red (Acer ginnala, Rhus typhnia, Parenthocissus quinquefolia, Viburnum dentatum) and four yellow species (Acer negundo, Ostrya virginiana, Vitis riparia, Zanthoxylum americanum) throughout autumn senescence and analyzed pigments, protein content, and chlorophyll fluorescence. We found yellow species retained the PsbS protein at higher levels, and had higher dark retention of zeaxanthin in late autumn relative to red species. All species retained lutein and the pool of xanthophyll cycle pigments in higher amounts than other carotenoids in late autumn. Our data support the hypothesis that red species use anthocyanins as a photoprotective strategy during autumn senescence, and therefore make less use of PsbS and zeaxanthin-mediated thermal dissipation. We also found species-specific variation in the particular combination of photoprotective strategies used.

Both external and internal factors induce heterogeneity in senescing leaves of deciduous trees.

Autumn senescence is characterised by spatial and temporal heterogeneity. We show that senescing birch (Betula spp.) leaves had lower PSII activity (probed by the F V /F M chlorophyll a fluorescence parameter) in late autumn than in early autumn. We confirmed that PSII repair slows down with decreasing temperature, while rates of photodamage and recovery, measured under laboratory conditions at 20°C, were similar in these leaves. We propose that low temperatures during late autumn hinder repair and lead to accumulation of non-functional PSII units in senescing leaves. Fluorescence imaging of birch revealed that chlorophyll preferentially disappeared from inter-veinal leaf areas. These areas showed no recovery capacity and low non-photochemical quenching while green veinal areas of senescing leaves resembled green leaves. However, green and yellow leaf areas showed similar values of photochemical quenching. Analyses of thylakoids isolated from maple (Acer platanoides ) leaves showed that red, senescing leaves contained high amounts of carotenoids and α-tocopherol, and our calculations suggest that α-tocopherol was synthesised during autumn. Thylakoids isolated from red maple leaves produced little singlet oxygen, probably due to the high antioxidant content. However, the rate of PSII photodamage did not decrease. The data show that the heterogeneity of senescing leaves must be taken into account to fully understand autumn senescence.

The clockwork of spring: bud dormancy timing as a driver of spring leaf-out in temperate deciduous trees

The initiation of spring leaf-out is a critical determinant of the growing season in trees, affecting primary production and species interactions in forest ecosystems. Variations in the timing of leaf-out among tree species are linked to their differential progression of bud dormancy. However, identifying reliable markers for bud dormancy has been challenging, leaving the connection between the timing of autumn leaf senescence, bud dormancy and spring leaf-out unclear. To test whether species initiating dormancy release earlier also exhibit earlier leaf senescence in autumn and earlier leaf-out in spring, we estimated the dates of peak dormancy depth (PDD), senescence timing and spring leaf-out across various species, locations and experimental conditions in Central Europe. PDD was defined as the date when the maximum thermal sum was required for leaf-out, whereas leaf senescence was assessed through the decrease in leaf greenness. Our findings reveal that PDD timing is a more accurate predictor of species-level differences in spring leaf-out dates than the timing of leaf senescence, the latter being a poor proxy for PDD. The observed temporal asynchrony between PDD and senescence was linked to dormancy induction showing species-specific sensitivity to temperature variations. Conversely, the timing of leaf senescence showed a consistent reaction to temperature changes across all species. These findings suggest that the physiological processes within buds and leaves during autumn are governed by distinct environmental cues, with the bud dormancy process serving as a more reliable predictor of spring phenological differences among forest tree species than does autumn leaf senescence.

Analysis of Nitrogen Dynamics and Transcriptomic Activity Revealed a Pivotal Role of Some Amino Acid Transporters in Nitrogen Remobilization in Poplar Senescing Leaves.

Leaf senescence is an important developmental process for deciduous trees during which part of leaf nitrogen is remobilized to branches, thus being beneficial for nitrogen conservation. However, the associated regulatory mechanism remains largely unknown in deciduous trees. In this study, nitrogen dynamics and transcriptomic activity in senescing leaves were measured during autumnal senescence in hybrid poplar. Both concentrations of leaf total nitrogen (N) and amine compounds were found to decline from the pre-senescence (PRE) to the middle-senescence (MS) stage. Although the total N concentration decreased further from MS to the late-senescence (LS) and leveled off to abscission (ABS) stage, amine compound concentration increased continuously from MS to ABS, suggesting that translocation of amine compounds underperformed production of amine compounds in leaves during this period. L-glutamate, L-glutamine and α-aminoadipic acid were the top three amine compounds accumulated in senescent leaves. RNA-Seq profiling identified thousands of differentially expressed genes (DEGs) with functional association with a metabolic transition towards disassimilation. Many genes encoding amino acid metabolism enzymes and amino acid transporters (AATs) were up-regulated. Comparison of expression trend with leaf N dynamics and phylogenetic analysis identified several PtAATs which exhibited down-regulation from MS to LS stage and putatively limited leaf N remobilization. This study can serve as a primary basis to further elucidate the molecular mechanisms of nitrogen remobilization in poplar senescing leaves.

Forage senescence and disease influence elk pregnancy across the Greater Yellowstone Ecosystem

AbstractFor various temperate ungulate species, recent research has highlighted the potential for spring vegetation phenology (“green‐up”) to influence individual condition, with purported benefits to population productivity. However, few studies have been able to measure the benefit on vital rates directly, and fewer still have investigated the comparative influence of other phenological periods on ungulate vital rates. In this study, we tracked phenological changes throughout the duration of the growing season and examined how their timing affected the probability of pregnancy in an ungulate population. We did this for elk (Cervus canadensis) across the Greater Yellowstone Ecosystem (GYE) by sampling 1106 adult females in winter at 25 sites over a 13‐year period and assessing sources of variation in pregnancy using a Bayesian hierarchical model. Pregnancy rates were generally high across the GYE (82.4%), and the primary influences on probability of pregnancy were the timing of vegetation senescence (“brown‐down”) in autumn and exposure to the reproductive disease brucellosis. Earlier forage brown‐down in fall negatively influenced the probability of pregnancy of elk aged 6–9 years by an estimated 17.2% within the range (ca. 32 days) of observed brown‐down end dates. While summer habitat quality has been inferred to influence elk pregnancy previously, our findings specify the key influence of foraging conditions later in the seasonal cycle, immediately before the breeding season. The reproductive disease brucellosis was also an important factor, reducing the probability of pregnancy by 12.4% in elk in the 6‐ to 9‐year age class. Because pregnancy was tested before most disease‐induced abortions occur, the apparent mechanism for this effect is a prolonged reduction in fertility beyond the period of initial exposure in which fetal mortality is typically expected. Our results prompt greater scrutiny of the combined effects of late‐season phenology and disease on reproductive rates and population productivity in temperate ungulates.

Characterization of autumn senescence in the Swedish Central Collection of heritage apple cultivars

Characterization of autumn senescence in the Swedish Central Collection of heritage apple cultivars

Biophysical and molecular characteristics of senescing leaves of two Norway maple varieties differing in anthocyanin content.

Disassembly and degradation of the photosynthetic protein complexes during autumn senescence, a vital step to ensure efficient nutrient relocalization for winter storage, is poorly understood. Concomitantly with the degradation, anthocyanins are often synthesized. However, as to why leaves accumulate red pigments, no consensus exists. One possibility is that anthocyanins protect senescing leaves from excess light. In this study, we investigated the pigment composition, photosynthetic performance, radical production, and degradation of the photosynthetic protein complexes in Norway maple (Acer platanoides) and in its highly pigmented, purple-colored variety (Faassen's black) during autumn senescence, to dissect the possible roles of anthocyanins in photoprotection. Our findings show that senescing Faassen's black was indeed more resistant to Photosystem II (PSII) photoinhibition, presumably due to its high anthocyanin content, than the green maple. However, senescing Faassen's black exhibited low photosynthetic performance, probably due to a poor capacity to repair PSII. Furthermore, an analysis of photosynthetic protein complexes demonstrated that in both maple varieties, the supercomplexes consisting of PSII and its antenna were disassembled first, followed by the degradation of the PSII core, Photosystem I, Cytochrome b6 f, and ATP synthase. Strikingly, the degradation process appeared to proceed faster in Faassen's black, possibly explaining its poor PSII repair capacity. The results suggest that tolerance against PSII photoinhibition may not necessarily translate to a better fitness. Finally, thylakoids isolated from senescing and non-senescing leaves of both maple varieties accumulated very little carbon-centered radicals, suggesting that thylakoids may not be a major source of reactive oxygen species in senescing leaves.

Salicylic acid metabolism and signalling coordinate senescence initiation in aspen in nature

Deciduous trees exhibit a spectacular phenomenon of autumn senescence driven by the seasonality of their growth environment, yet there is no consensus which external or internal cues trigger it. Senescence starts at different times in European aspen (Populus tremula L.) genotypes grown in same location. By integrating omics studies, we demonstrate that aspen genotypes utilize similar transcriptional cascades and metabolic cues to initiate senescence, but at different times during autumn. The timing of autumn senescence initiation appeared to be controlled by two consecutive “switches”; 1) first the environmental variation induced the rewiring of the transcriptional network, stress signalling pathways and metabolic perturbations and 2) the start of senescence process was defined by the ability of the genotype to activate and sustain stress tolerance mechanisms mediated by salicylic acid. We propose that salicylic acid represses the onset of leaf senescence in stressful natural conditions, rather than promoting it as often observed in annual plants.

Identification of ApbHLH1 as a Partner Interacting with ApMYB1 to Promote Anthocyanin Biosynthesis during Autumnal Leaf Coloration in Acer palmatum

Anthocyanin biosynthesis determines the leaf color of Acer palmatum as a widely-planted landscape tree. Previously, ApMYB1 has been characterized as a positive regulator of anthocyanin biosynthesis. To further elucidate the mechanism of leaf coloration, the present study identified a basic helix-loop-helix (bHLH) transcription factor (ApbHLH1) through the phylogenetic analysis of 156 putative bHLH proteins in Acer palmatum and eight reference bHLHs which were known to be involved in the anthocyanin biosynthesis of selected plants. Protein structure comparison showed that ApbHLH1 has a conserved bHLH domain, and its N-terminal contains an MYB-interacting region. The expression of ApbHLH1 in leaves was found to not be correlated with anthocyanin contents either in green, semi-red leaves or during leaf autumnal senescence when anthocyanin content increased. ApbHLH1 expression in detached leaves was induced by exogenous senescence-promoting chemicals, including H2O2, SA, MeJA, ACC and ABA, with certain durations. In particular, either high light or low temperature induced ApbHLH1 expression significantly, and combination of high light and low temperatures seemed more effective in inducing ApbHLH1 expression. Luciferase complementation imaging assays confirmed the physical interaction between ApbHLH1 and ApMYB1, which could be abolished by either the truncating MYB-interacting region of ApbHLH1 or the deleting bHLH interacting domain of ApMYB1. The transient expression of ApbHLH1 could not induce anthocyanin production, while the co-expression of ApbHLH1 and ApMYB1 resulted in a higher accumulation of anthocyanins compared to the expression of ApMYB1 alone in tobacco leaves. Collectively, our results revealed that ApbHLH1 participated in leaf coloration through binding with ApMYB1 and enhancing the ApMYB1 function of promoting anthocyanin biosynthesis during leaf autumnal reddening in Acer palmatum. ApbHLH1 could have the potential for breeding color-leafed plants through co-transformation with ApMYB1.

Autumn canopy senescence has slowed down with global warming since the 1980s in the Northern Hemisphere

Climate change strongly impact vegetation phenology, with considerable potential to alter land-atmosphere carbon dioxide exchange and terrestrial carbon cycle. In contrast to well-studied spring leaf-out, the timing and magnitude of autumn senescence remains poorly understood. Here, we use monthly decreases in Normalized Difference Vegetation Index satellite retrievals and their trends to surrogate the speed of autumn senescence during 1982–2018 in the Northern Hemisphere (>30°N). We find that climate warming accelerated senescence in July, but this influence usually reversed in later summer and early autumn. Interestingly, summer greening causes canopy senescence to appear later compared to an advancing trend after eliminating the greening effect. This finding suggests that summer canopy greening may counteract the intrinsic changes in autumnal leaf senescence. Our analysis of autumn vegetation behavior provides reliable guidance for developing and parameterizing land surface models that contain an interactive dynamic vegetation module for placement in coupled Earth System Models.

Modeling seasonal vegetation phenology from hydroclimatic drivers for contrasting plant functional groups within drylands of the Southwestern USA

In dryland ecosystems, vegetation within different plant functional groups exhibits distinct seasonal phenologies that are affected by the prevailing hydroclimatic forcing. The seasonal variability of precipitation, atmospheric evaporative demand, and streamflow influences root-zone water availability to plants in water-limited environments. Increasing interannual variations in climate forcing of the local water balance and uncertainty regarding climate change projections have raised the potential for phenological shifts and changes to vegetation dynamics. This poses significant risks to plant functional types across large areas, especially in drylands and within riparian ecosystems. Due to the complex interactions between climate, water availability, and seasonal plant water use, the timing and amplitude of phenological responses to specific hydroclimate forcing cannot be determined a priori, thus limiting efforts to dynamically predict vegetation greenness under future climate change. Here, we analyze two decades (1994–2021) of remote sensing data (soil adjusted vegetation index (SAVI)) as well as contemporaneous hydroclimate data (precipitation, potential evapotranspiration, depth to groundwater, and air temperature), to identify and quantify the key hydroclimatic controls on the timing and amplitude of seasonal greenness. We focus on key phenological events across four different plant functional groups occupying distinct locations and rooting depths in dryland SE Arizona: semi-arid grasses and shrubs, xeric riparian terrace and hydric riparian floodplain trees. We find that key phenological events such as spring and summer greenness peaks in grass and shrubs are strongly driven by contributions from antecedent spring and monsoonal precipitation, respectively. Meanwhile seasonal canopy greenness in floodplain and terrace vegetation showed strong response to groundwater depth as well as antecedent available precipitation (aaP = P − PET) throughout reaches of perennial and intermediate streamflow permanence. The timings of spring green-up and autumn senescence were driven by seasonal changes in air temperature for all plant functional groups. Based on these findings, we develop and test a simple, empirical phenology model, that predicts the timing and amplitude of greenness based on hydroclimate forcing. We demonstrate the feasibility of the model by exploring simple, plausible climate change scenarios, which may inform our understanding of phenological shifts in dryland plant communities and may ultimately improve our predictive capability of investigating and predicting climate-phenology interactions in the future.

Not all temperate deciduous trees are leafless in winter: The curious case of marcescence

AbstractTemperate deciduous forests by definition include a large proportion of woody species that shed their leaves each autumn and are completely leafless during winter months. Leaf senescence in deciduous trees is an active, complex process typically involving the physiological formation of an abscission layer causing the petiole to mechanically detach from the branch. However, several deciduous species retain all or some senesced leaves on branches through much of winter, a phenomenon called leaf marcescence. Marcescence has long fascinated botanists, including Pehr Kalm as early as 1749. Yet, surprisingly little research has been done to date. Here, we review and explore patterns and mechanisms of leaf marcescence in temperate forests, bringing together six nonmutually exclusive but separately proposed hypotheses: (1) Marcescence has no adaptive function but rather an evolutionary byproduct; (2) Marcescent leaves deter winter browsing herbivores; (3) Leaf retention through winter improves nutrient resorption during autumn senescence; (4) Prolonged leaf shedding into spring minimizes nutrient leaching and promotes decomposition; (5) Marcescent leaves protect overwintering buds from frost or desiccation; and (6) Marcescent canopies provide winter cover for animals (including insects, birds, bats), thereby affecting plant nutrient availability via excrement. No hypothesis has complete support and few tests of multiple hypotheses have been done. It is likely that any adaptive value of marcescence is species and context dependent. Despite increased interest in plant phenology and prevalence of this trait, much remains to be understood on the physiology, evolution, function, and ecological implications of leaf marcescence.

Increased precipitation leads to earlier green-up and later senescence in Tibetan alpine grassland regardless of warming

It is debatable whether warming or increased precipitation primarily drives the changes of spring and autumn phenology in alpine grasslands at high elevations like the Tibetan Plateau. We aim to test the hypothesis that increased precipitation and soil moisture rather than warming significantly advance spring green-up dates (GUD) of dominant species in a semiarid alpine grassland, while both increases of temperature and precipitation delay their autumn senescence dates (SD). We conducted a 2-year manipulative experiment with infrared warming (ambient, +2 °C) and precipitation increase for each of rainfall events (ambient, +15 %, +30 %) during the growing season in a Tibetan alpine grassland. GUD and SD of three dominant species and the relevant soil temperature (ST) and moisture (SM) were observed. Rainy season onset as well as Pre-GUD or Pre-SD (30 days before GUD or SD) mean air-temperature (T-30d) and precipitation (P-30d) and relevant soil temperature (ST-30d) and moisture (SM-30d) were calculated for each experimental treatment. GUD dates of the three dominant species were advanced by increased precipitation rather than by warming, which showed a robust positive correlation with rainy season onset. SD dates were independently delayed by both increases of temperature and precipitation. There was no interactive effect of warming and increased precipitation on GUD and SD across species and years. In general, GUD had a significant negative correlation with Pre-GUD P-30d (SM-30d) but not with Pre-GUD T-30d (ST-30d), while SD showed a significant positive correlation with Pre-SD T-30d and P-30d or Pre-SD ST-30d and SM-30d. Our data support the hypothesis, indicating that spring and autumn phenology of monsoon-adapted alpine vegetation are more sensitive to precipitation change than to warming. The prolonged growing season length under increased temperature and precipitation is more depended on the delay of autumn senescence than the advance of spring green-up.

Red pigments in autumn leaves of Norway maple do not offer significant photoprotection but coincide with stress symptoms.

The reasons behind autumn colors, a striking manifestation of anthocyanin synthesis in plants, are poorly understood. Usually, not all leaves of an anthocyanic plant turn red or only a part of the leaf blade turns red. In the present study, we compared green, red and yellow sections of senescing Norway maple leaves, asking if red pigments offer photoprotection, and if so, whether the protection benefits the senescing tree. Green and senescing maple leaves were illuminated with strong white, green or red light in the absence or presence of lincomycin which blocks photosystem II (PSII) repair. Irrespective of the presence of anthocyanins, senescing leaves showed weaker capacity to repair PSII than green leaves. Furthermore, the rate of photoinhibition of PSII did not significantly differ between red and yellow sections of senescing maple leaves. We also followed pigment contents and photosynthetic reactions in individual leaves, from the end of summer until abscission of the leaf. In maple, red pigments accumulated only during late senescence, but light reactions stayed active until most of the chlorophyll had been degraded. PSII activity was found to be lower and non-photochemical quenching higher in red leaf sections, compared with yellow sections of senescing leaves. Red leaf sections were also thicker. We suggest that the primary function of anthocyanin synthesis is not to protect senescing leaves from excess light but to dispose of carbohydrates. This would relieve photosynthetic control, allowing the light reactions to produce energy for nutrient translocation at the last phase of autumn senescence when carbon skeletons are no longer needed.

Multi-Temporal Assessment of Remotely Sensed Autumn Grass Senescence across Climatic and Topographic Gradients

Climate and topography are influential variables in the autumn senescence of grassland ecosystems. For instance, extreme weather can lead to earlier or later senescence than normal, while higher altitudes often favor early grass senescence. However, to date, there is no comprehensive understanding of key remote-sensing-derived environmental variables that influence the occurrence of autumn grassland senescence, particularly in tropical and subtropical regions. Meanwhile, knowledge of the relationship between autumn grass senescence and environmental variables is required to aid the formulation of optimal rangeland management practices. Therefore, this study aimed to examine the spatial autocorrelations between remotely sensed autumn grass senescence vis-a-vis climatic and topographic variables in the subtropical grasslands. Sentinel 2′s Normalized Difference NIR/Rededge Normalized Difference Red-Edge (NDRE) and the Chlorophyll Red-Edge (Chlred-edge) indices were used as best proxies to explain the occurrence of autumn grassland senescence, while monthly (i.e., March to June) estimates of the remotely sensed autumn grass senescence were examined against their corresponding climatic and topographic factors using the Partial Least Square Regression (PLSR), the Multiple Linear Regression (MLR), the Classification and Regression Trees (CART), and the Random Forest Regression (RFR) models. The RFR model displayed a superior performance on both proxies (i.e., RMSEs of 0.017, 0.012, 0.056, and 0.013, as well as R2s of 0.69, 0.71, 0.56, and 0.71 for the NDRE, with RMSEs and R2s 0.023, 0.018, 0.014 and 0.056, as well as 0.59, 0.60, 0.69, and 0.72 for the Chlred-edge in March, April, May, and June, respectively). Next, the mean monthly values of the remotely sensed autumn grass senescence were separately tested for significance against the average monthly climatic (i.e., minimum (Tmin) and maximum (Tmax) air temperatures, rainfall, soil moisture, and solar radiation) and topographic (i.e., slope, aspect, and elevation) factors to define the environmental drivers of autumn grassland senescence. Overall, the results indicated that Tmax (p = 0.000 and 0.005 for the NDRE and the Chlred-edge, respectively), Tmin (p = 0.021 and 0.041 for the NDRE and the Chlred-edge, respectively), and the soil moisture (p = 0.031 and 0.040 for the NDRE and the Chlred-edge, respectively) were the most influential autumn grass senescence drivers. Overall, these results have shown the role of remote sensing techniques in assessing autumn grassland senescence along climatic and topographic gradients as well as in determining key environmental drivers of this senescence in the study area