Interactive effects of soil nitrogen and water availability on leaf mass loss in a temperate steppe

Although the link between leaf mass loss and assessment of ecosystem nutrient use efficiency and plant nutrient resorption efficiency has received considerable attention in various ecosystems, there has been relatively little effort to assess plant leaf mass loss during senescence, especially for herbaceous species. We conducted experimental studies to assess leaf mass loss during senescence in five dominant herbaceous species and examined the effects of increasing nitrogen (N) and water availability on leaf mass loss of four species in a temperate steppe in northern China. We nondestructively estimated mature leaf mass based on leaf length and width. Leaf mass loss varied substantially among species, ranging from 20–50%. On average across all species, N and water addition increased leaf mass loss by 30% and 19%, respectively. N and water addition interacted to affect leaf mass loss, as water addition had a significant positive effect on leaf mass loss under enriched N conditions but showed no effect under ambient N levels. We conclude that leaf mass loss of herbaceous plants was considerable and can potentially be more pronounced with increasing N and water availability. It is notable that the responses of plant species to N and water addition were variable. We suggest that leaf mass loss during senescence should be given full consideration in assessing nutrient use and resorption efficiency in semi-arid areas.     

Does the Gradualness of Leaf Shedding Govern Nutrient Resorption from Senescing Leaves in Mediterranean Woody Plants?

To reveal the environmental and substrate quality effects on decomposition process and enzyme activities, litterbag experiments containing Nuphar and Carex leaves, Nuphar rhizome, and Ranunculus shoot, were carried in five-subalpine marshes in Lake Tahoe basin, USA. Alkaline phosphatase, β-glucosidase, and β-xylosidase activities were determined by a fluorogenic method using methyumbelliferyl substrates. Carex leaves, Nuphar rhizome and leaves, and Ranunculus shoots lost, respectively, 33, 67, 82 and 93% of original dry weight over 268 days. Decay rates were different among substrates but not among marshes. Nitrogen and carbon contents increased during the first 58 days and subsequently remained stable. Phosphorus content was stable during the experimental period except for a decrease in the first 16 days in Nuphar shoots. Enzyme activities in decomposing Carex and Nuphar leaves in four marshes were not significantly affected by environmental conditions. β-glucosidase and β-xylosidase activities in decomposing Carex leaves increased with time, but in other plant tissue these enzyme activities remained stable during experimental period. Enzyme activities were significantly different among decomposing substrates. Alkaline phosphatase activity was highest in Nuphar leaves (ca. 1286 μ-mole h⁻¹ g DW −1) but lower and similar in other plant tissues (ca. 100 and 10 μ-mole h −1 g DW −1, respectively). This study showed differences in decay rates and enzyme activities rely on substrate and not the environment conditions of the study area. Decomposition rates in the early stage of decomposition were related to cumulative enzyme activities.     

Seasonal changes in element contents in mangrove element retranslocation during leaf senescene

The concentrations of nitrogen (N), phosphorus (P), potassium (K), sodium (Na), calcium (Ca), magnesium (Mg) and chlorine (Cl) were followed monthly in pre-senescence leaves and post-abscission leaves of Kandelia candel (L.) Druce at the Jiulongjiang estuary, and Fujian, China. The element retranslocation efficiency (RE) was studied during leaf senescence. The element RE's evaluated using different methods were compared and a new method was put forward to evaluate element RE during leaf senescence in evergreen trees without concentrated leaf fall. The results showed that during leaf senescence, 77.22% N, 57.53% P, and 44.51% K were translocated out of senescing leaves. Translocation of nutrients out of senescing leaves back into shoots was an important nutnent-conservation mechanism for N and P, was less important for K, and did not occur for Ca, Mg, Na, or Cl. One of the reasons for the high primary productivity of mangroves in nutrient poor sites (especially with low N) is the high nutrient use efficiency.     

Drought-induced senescence is characterized by a loss of antioxidant defences in chloroplasts

The relationship between drought, oxidative stress and leaf senescence was evaluated in field‐grown sage (Salvia officinalis L.), a drought‐susceptible species that shows symptoms of senescence when exposed to stress. Despite the photoprotection conferred by the xanthophyll cycle, drought‐stressed senescing leaves showed enhanced lipid peroxidation, chlorophyll loss, reduced photosynthetic activity and strong reductions of membrane‐bound chloroplastic antioxidant defences (i.e. β‐carotene and α‐tocopherol), which is indicative of oxidative stress in chloroplasts. H₂O₂ accumulated in drought‐stressed senescing leaves. Subcellular localization studies showed that H₂O₂ accumulated first in xylem vessels and the cell wall and later in the plasma membrane of mesophyll cells, but not in chloroplasts, indicating reactive oxygen species other than H₂O₂ as direct responsible for the oxidative stress observed in the chloroplasts of drought‐stressed senescing leaves. The strong degradation of β‐carotene and α‐tocopherol suggests an enhanced formation of singlet oxygen as the putative reactive oxygen species responsible for oxidative stress to senescing chloroplasts. This study demonstrates that oxidative stress in chloroplasts mediates drought‐induced leaf senescence in sage growing in Mediterranean field conditions.     

Drought-deciduous behavior reduces nutrient losses from temperate deciduous trees under severe drought

Nutrient resorption from senescing leaves is an important mechanism of nutrient conservation in temperate deciduous forests. Resorption, however, may be curtailed by climatic events that cause rapid leaf death, such as severe drought, which has been projected to double by the year 2100 in the eastern United States. During a record drought in the southeastern US, we studied 18 common temperate winter-deciduous trees and shrubs to understand how extreme drought affects nutrient resorption of the macronutrients N, P, K, and Ca. Four species exhibited drought-induced leaf senescence and maintained higher leaf water potentials than the remaining 14 species (here called drought-evergreen species). This strategy prevented extensive leaf desiccation during the drought and successfully averted large nutrient losses caused by leaf desiccation. These four drought-deciduous species were also able to resorb N, P, and K from drought-senesced leaves, whereas drought-evergreen species did not resorb any nutrients from leaves lost to desiccation during the drought. For Oxydendrum arboreum, the species most severely affected by the drought, our results indicate that trees lost 50% more N and P due to desiccation than would have been lost from fall senescence alone. For all drought-deciduous species, resorption of N and P in fall-senesced leaves was highly proficient, whereas resorption was incomplete for drought-evergreen species. The lower seasonal nutrient losses of drought-deciduous species may give them a competitive advantage over drought-evergreen species in the years following the drought, thereby impacting species composition in temperate deciduous forests in the future.     

Extended leaf senescence promotes carbon gain and nutrient resorption: importance of maintaining winter photosynthesis in subtropical forests

The relative advantages of being deciduous or evergreen in subtropical forests and the relationship between leaf phenology and nutrient resorption efficiency are not well understood. The most successful deciduous species (Lyonia ovalifolia) in an evergreen-dominated subtropical montane cloud forest in southwest (SW) China maintains red senescing leaves throughout much of the winter. The aim of this study was to investigate whether red senescing leaves of this species were able to assimilate carbon in winter, to infer the importance of maintaining a positive winter carbon balance in subtropical forests, and to test whether an extended leaf life span is associated with enhanced nutrient resorption and yearly carbon gain. The red senescing leaves of L. ovalifolia assimilated considerable carbon during part of the winter, resulting in a higher yearly carbon gain than co-occurring deciduous species. Its leaf N and P resorption efficiency was higher than for co-occurring non-anthocyanic deciduous species that dropped leaves in autumn, supporting the hypothesis that anthocyanin accumulation and/or extended leaf senescence help in nutrient resorption. Substantial winter carbon gain and efficient nutrient resorption may partially explain the success of L. ovalifolia versus that of the other deciduous species in this subtropical forest. The importance of maintaining a positive carbon balance for ecological success in this forest also provides indirect evidence for the dominance of evergreen species in the subtropical forests of SW China.     

The efficiency of nitrogen retranslocation from leaf biomass in Quercus ilex ecosystems

Nitrogen retranslocation from senescing leaves represents a crucial adaptation by tree species towards a more efficient use of this nutrient. As a result, this part of the nitrogen cycle has received increasing attention in recent years. However, there remain strong discrepancies with respect to the factors responsible for interspecific differences in the efficiency of this process.In the present work the seasonal pattern of leaf growth and the movement of nitrogen in leaves have been studied in a series of Quercus ilex plots with different levels of rainfall and soil quality in central-western Spain, as well as in 20 other woody species typical of this area. The percentage of nitrogen retranslocated was estimated from the difference between the maximum mass of nitrogen stored in the leaf biomass and the amount of this nutrient returned annually to the soil through leaf fall. Q. ilex appears as one of the least efficient species in the Mediterranean region in the recovery of nitrogen from senescing leaves (29.7% of the maximum pool). Furthermore, the older leaves of Q. ilex do not show the cycles of nitrogen withdrawal during new flushes of shoot growth, such as occurs in Pinus spp. This suggests that older leaves in Q. ilex do not play an important role as nitrogen storage organs.     

Ammonium and proline accumulation in senescing cut leaves of <Emphasis Type="Italic">Zantedeschia</Emphasis>

Accumulation of ammonium and proline were reported as phenomena associated with plant response to stress and/or senescence. The effects of a preservative (8HQC + sucrose) and 24 hrs pulse conditioning with GA₃ on the ammonium and proline contents were studied in senescing cut leaves of Zantedeschia aethiopica Spr. and Z. elliottiana Engl., grown for the florists green. Generally, accumulation of both compounds was observed in senescing leaves, however, the final ammonium and proline levels depended upon the species and the treatment applied. Conditioning with GA₃, a treatment known to delay leaf senescence in Zantedeschia sp., prevented the increases in the ammonium and proline contents. Standard preservative solution used to prolong the longevity of cut flowers enhanced the ammonium accumulation in senescing leaves of both species, and the proline accumulation in the leaves of Z. aethiopica, but not in Z. elliottiana. These observations suggest that neither ammonium nor proline accumulation would be fully reliable predictors of cut leaf freshness during their entire market life. However, proline accumulation could serve as a quick test of freshness in the first half of the useful market life of cut leaves of Zantedeschia.     

Current measures of nutrient resorption efficiency lead to a substantial underestimation of real resorption efficiency: facts and solutions

Nutrient resorption is an important process during leaf senescence, which helps plants to minimize nutrient losses. To quantify nutrient resorption, the parameter resorption efficiency is commonly used. This parameter describes the percentage of the nutrient pool withdrawn before leaf abscission. The nutrient pool is generally expressed on the basis of leaf mass or leaf area, assuming that these bases do not change during senescence. In this paper we firstly present a mathematical formula describing the effect of change in measurement basis on the difference between the real resorption efficiency (RRE) value and the measured resorption efficiency (MRE). This formula shows that even moderate senescence-related changes in a measurement basis can lead to considerable underestimation of RRE. Secondly, to estimate the general change in measurement basis we quantified leaf mass loss and leaf shrinkage during senescence from literature data. These data shows that mass loss percentages can be as high as 40%, and leaf shrinkage can be up to 20%. This level of change in basis seriously compromises the MRE when not corrected for. Using our formula and the reported average literature values of changes in leaf mass (21%) and leaf shrinkage (11%) during senescence, we calculated that the average RRE for nitrogen and phosphorous of terrestrial plants is 6% (leaf area) to 10% (leaf mass) higher than the 50%, respectively 52% as reported by Aerts (1996) . This implies that nutrient resorption from senescing leaves is even more important for nutrient retention in terrestrial plants than thought so far. We advocate that preselecting leaves and monitoring the measurement basis throughout the duration of the experiment should minimize the difference between MRE and RRE.     

Delayed leaf senescence in ethylene-deficient ACC-oxidase antisense tomato plants: molecular and physiological analysis

To determine the role of ethylene during tomato (Lycopersicon esculentum Mill. cv. Alisa Craig) leaf senescence, transgenic ACC oxidase antisense plants were analysed. Northern analysis of wild-type plants indicated that ACC oxidase mRNA accumulation normally begins in pre-senescent green leaves but was severely reduced in the antisense plants. Although the levels of ethylene evolved by wild-type and transgenic leaves increased during the progression of senescence, levels were extremely low in transgenic leaves. Leaf senescence, as assessed by colour change from green to yellow, was clearly delayed by 10–14 days in the antisense plants when compared with wild-type plants. Northern analysis of the photosynthesis-associated genes, cab and rbcS, indicated that levels of the corresponding mRNAs were higher in transgenic leaves which were not yet senescing compared with senescing wild-type leaves of exactly the same age. Northern analysis using probes for tomato fruit ripening-related genes expressed during leaf senescence indicated that once senescence was initiated the expression pattern of these mRNAs was similar in transgenic and wild-type leaves. In the antisense plants chlorophyll levels, photosynthetic capacity and chlorophyll fluorescence were higher when compared with senescing wild-type plants of the same age. Photosynthetic capacity and the quantum efficiency of photosystem II were maintained for longer in the transformed plants at values close to those observed in wild-type leaves prior to the visible onset of senescence. These results indicate that inhibiting ACC oxidase expression and ethylene synthesis results in delayed leaf senescence, rather than inducing a stay-green phenotype. Once senescence begins, it progresses normally. Onset of senescence is not, therefore, related to a critical level of ethylene. The correlation between higher levels prior to senescence and early onset, however, suggests that ethylene experienced by the plant may be a significant contributing factor in the timing of senescence.     

Mass loss,fungal colonisation and nutrient dynamics of Phragmites australis leaves during senescence and early aerial decay

This study examined the mass loss, fungal biomass, and nutrient dynamics of standing Phragmites australis leaf blades during senescence and early decay in littoral reed stands of two hardwater lakes. Green living leaves were tagged at defined canopy heights in early autumn (late August or early September) and periodically collected until all leaf blades had fallen off the parent shoot. Samples were analysed for leaf dry mass remaining, fungal biomass associated with leaves (ergosterol concentrations), and nitrogen and phosphorus concentrations. Considerable mass loss of leaves occurred in the standing position (up to 28%). Nitrogen and phosphorus concentrations of leaves decreased substantially with time (by 39–77%), indicating that a major portion of these nutrients was translocated to the rhizome during senescence. Fungal biomass associated with leaves increased during the study period, reaching an estimated maximum of about 40 mg g⁻¹ of leaf dry mass. Fungal biomass was negatively correlated with leaf N and P concentrations. The observed patterns of leaf mass loss, nutrient dynamics, and fungal biomass were consistent with the successive senescence and death of leaves from the shoot base to its tip. The results of this study point to a notable mass loss of P. australis leaf blades in the standing position, which appears to be mediated by both plant and microbial processes. Nutrient dynamics, in contrast, appear to be largely governed by plant processes.     

Water- and nutrient-use efficiency of a deciduous species, Vaccinium myrtillus, and an evergreen species, V. vitis-idaea, in a subalpine dwarf shrub heath in the southern Alps, Italy

Periodic measurements of gas‐exchange rates and determinations of foliar N and P concentrations were used for evaluating instantaneous water‐use efficiency and photosynthetic nutrient‐use efficiency in two co‐existing dwarf shrubs of different growth form (V. myrtillus, deciduous, and V. vitis‐idaea, evergreen) in a subalpine heath in the southern Alps of Italy. Those data were compared with cumulative assessments of water‐use efficiency and photosynthetic nutrient‐use efficiency obtained by measuring leaf carbon isotope discrimination in leaf tissues and by estimating nutrient resorption from senescing leaves. V. myrtillus presented higher dry‐weight based rates of net photosynthesis (A_weight) compared to V. vitis‐idaea. A_weight was positively correlated with foliar‐nutrient status and intercellular‐to‐ambient gradient in CO₂ concentrations. A_weight was, furthermore, negatively correlated with leaf specific mass. Instantaneous photosynthetic nutrient‐use efficiency did not differ between the two species but the percentages of N and P pools resorbed from senescing leaves were somewhat higher in the deciduous species. The evergreen species showed lower P concentrations in senescing leaves which indicated a higher proficiency in resorbing phosphorus compared to the deciduous species. In addition, the evergreen species achieved a higher carbon gain per unit foliar N and P, due to a longer mean residence time of both nutrients. The two species did not differ from each other with respect to both instantaneous and long‐term water‐use efficiency. This was consistent with the climatic pattern, showing no sign of water deficiency through the growing season. Current‐year V. vitis‐idaea leaves had a significantly higher Δ¹³C compared to previous‐year leaves, possibly mirroring a long term acclimation of evergreen leaves, as far as they age, to the habitat conditions in the understory where evergreen species are usually confined within mixed dwarf‐shrub communities.     

Possible mechanisms of light-induced chlorophyll degradation in senescing leaves of Hydrilla verticillata

Light treatment markedly accelerated the chlorophyll loss in senescing leaves of Hydrilla verticillata [(L.f.) Royle] as compared to dark treatment, whereas such acceleration could not be observed in senescing spinach (Spinacia oleracea L.) leaves. The light-induced cholorophyll loss in Hydrilla was retarded slightly by chloramphenicol and markedly by cycloheximide. Catalase (EC 1.11.1.6) activity did not change appreciably in Hydrilla leaves either in light or in darkness, while in spinach it declined markedly in the dark, and light retarded such decline. Peroxidase activity in Hydrilla showed faster increase in light than in darkness, while in spinach it increased only in light during senescence. The activity of phenol(pyrogallol)-specific peroxidase increased markedly in light, and that of ascorbate-specific peroxidase decreased slightly both in light and darkness during senescence of Hydrilla leaves. This rise in phenolspecific peroxidase activity was prevented by cycloheximide treatment. Pretreatment of Hydrilla leaves with monophenol (2,4-dichlorophenol) and o-diphenol (hydroquinone) accelerated and retarded, respectively, the light-induced cholorophyll loss. Pretreatment of Hydrilla leaves with H₂O₂ augmented the chlorophyll loss more markedly in light than in darkness. The endogenous level of H₂O₂ increased more in light than in dark during senescence of Hydrilla leaves. Treatment of Hydrilla leaves with 3-(3.4-dichlorophenyl)-l,l-dimethylurea. a photosystem II inhibitor, prevented both light-induced rise in H₂O: level and chlorophyll loss, but it was without effect in the dark. Retardation of light-induced chlorophyll loss occurred during senescence of Hydrilla leaves when light was given in different photoperiods in a 24-h daily cycle for 6 days instead of as continuous irradiance. There was a negative correlation between the length of the photoperiod and the extent of cholorophyll loss.     

Foliar demand and resource economy of nutrients in dry tropical forest species

Important phenological activities in seasonally dry tropical forest species occur within the hot‐dry period when soil water is limiting, while the subsequent wet period is utilized for carbon accumulation. Leaf emergence and leaf area expansion in most of these tree species precedes the rainy season when the weather is very dry and hot and the soil cannot support nutrient uptake by the plants. The nutrient requirement for leaf expansion during the dry summer period, however, is substantial in these species. We tested the hypothesis that the nutrients withdrawn from the senescing leaves support the emergence and expansion of leaves in dry tropical woody species to a significant extent. We examined the leaf traits (with parameters such as leaf life span, leaf nutrient content and retranslocation of nutrients during senescence) in eight selected tree species in northern India. The concentrations of N, P and K declined in the senescing foliage while those of Na and Ca increased. Time series observations on foliar nutrients indicated a substantial amount of nutrient resorption before senescence and a ‘tight nutrient budgeting’. The resorbed N‐mass could potentially support 50 to 100% and 46 to 80% of the leaf growth in terms of area and weight, respectively, across the eight species studied. Corresponding values for P were 29 to 100% and 20 to 91%, for K 29 to 100% and 20 to 57%, for Na 3 to 100% and 1 to 54%, and for Ca 0 to 32% and 0 to 30%. The species differed significantly with respect to their efficiency in nutrient resorption. Such interspecific differences in leaf nutrient economy enhance the conservative utilization of soil nutrients by the dry forest community. This reflects an adaptational strategy of the species growing on seasonally dry, nutrient‐poor soils as they tend to depend more or less on efficient internal cycling and, thus, utilize the retranslocated nutrients for the production of new foliage biomass in summer when the availability of soil moisture and nutrients is severely limited.     

Increased cytokinin levels in transgenic PSAG12–IPT tobacco plants have large direct and indirect effects on leaf senescence, photosynthesis and N partitioning

We studied the impact of delayed leaf senescence on the functioning of plants growing under conditions of nitrogen remobilization. Interactions between cytokinin metabolism, Rubisco and protein levels, photosynthesis and plant nitrogen partitioning were studied in transgenic tobacco (Nicotiana tabacum L.) plants showing delayed leaf senescence through a novel type of enhanced cytokinin syn‐thesis, i.e. targeted to senescing leaves and negatively auto‐regulated (P_SAG12–IPT), thus preventing developmental abnormalities. Plants were grown with growth‐limiting nitrogen supply. Compared to the wild‐type, endogenous levels of free zeatin (Z)‐ and Z riboside (ZR)‐type cytokinins were increased up to 15‐fold (total ZR up to 100‐fold) in senescing leaves, and twofold in younger leaves of P_SAG12–IPT. In these plants, the senescence‐associated declines in N, protein and Rubisco levels and photosynthesis rates were delayed. Senescing leaves accumulated more (¹⁵N‐labelled) N than younger leaves, associated with reduced shoot N accumulation (–60%) and a partially inverted canopy N profile in P_SAG12–IPT plants. While root N accumulation was not affected, N translocation to non‐senescing leaves was progressively reduced. We discuss potential consequences of these modified sink–source relations, associated with delayed leaf senescence, for plant productivity and the efficiency of utilization of light and minerals.     

Proteomic analysis of residual proteins in blades and petioles of fallen leaves of Brassica napus

Brassica napus L. is an important crop plant, characterised by high nitrogen (N) levels in fallen leaves, leading to a significant restitution of this element to the soil, with important consequences at the economic and environmental levels. It is now well established that the N in fallen leaves is due to weak N remobilisation that is especially related to incomplete degradation of foliar proteins during leaf senescence. Identification of residual proteins in a fallen leaf (i.e. incompletely degraded in the last step of the N remobilisation process) constitutes important information for improving nutrient use efficiency. Proteome analysis of the vascular system (petioles) and blades from fallen leaves of Brassica napus was performed, and the 30 most abundant residual proteins in each tissue were identified. Among them, several proteins involved in N recycling remain in the leaf after abscission. Moreover, this study reveals that some residual proteins are associated with energy metabolism, protection against oxidative stress, and more surprisingly, photosynthesis. Finally, comparison of blade and petiole proteomes show that, despite their different physiological roles in the non‐senescing leaf, both organs redirect their metabolism in order to ensure catabolic reactions. Taken together, the results suggest that a better degradation of these leaf proteins during the senescence process could enable improvements in the N use efficiency of Brassica napus.     

Foliar Nitrogen Dynamics and Nitrogen Resorption of a Sandy Shrub Salix gordejevii in Northern China

Resorption of nitrogen (N) from senescing leaves is an important conservation mechanism that allows plants to use the same N repeatedly. Seasonal variations in leaf nitrogen of mature green and senescing leaves and N resorption in Salix gordejevii Chang, a sandy shrub in northern China, were studied. Our objective was to compare N resorption of this Salix species that successfully occupy different habitats (shifting sandland, fixed sandland and lowland) with differences in soil N availability and moisture. Nitrogen concentrations in green and senescing leaves were higher in June and July. N resorption efficiency (percentage reduction of N between green and senescing leaves) was highest at shifting sandland, intermediate at fixed sandland, and lowest at lowland. There was a clear seasonal variation in N-resorption efficiency, with a lower value at the early growing season and a higher value during summer. N resorption efficiency was lower at the sites with higher soil N availability, suggesting that the efficiency of the resorption process is determined by the availability of the nutrient in the soil. Resorption from senescing leaves may play an important role in the nitrogen dynamics of sandy plants and reduce the nitrogen requirements for plant growth. We conclude that N resorption from senescing leaves in S. gordejevii was correlated to soil characteristics and higher N resorption on poor soils is a phenotypic adjustment by this species to maximize N-use at low availability.     

Qualitative and quantitative changes in nitrogenous compounds in senescing leaf and bark tissues of the apple

Quantitative and qualitative changes in proteins and ethanol-soluble nitrogen were followed in senescing leaf and bark tissues of ‘Golden Delicious’ apple trees (Malus domestica Borkh.). While senescing leaves lost 46% of their proteins, total bark protein increased 240% during senescence. However, the protein nitrogen gain in bark tissue was about the same as the protein nitrogen loss in leaf tissue per unit fresh weight of tissues. The pattern of bark protein accumulation appears to be gradual from early August to November and sequential from lower to higher molecular weight species of proteins. The final electrophoretic profile of total bark proteins was established at the later stages of senescence. By late November, 89% of the nitrogen in the bark tissue was found in proteins with 11% in the ethanol-soluble fractions. The total protein content of dormant bark tissue was 3.5% per gram dry tissue. Fractionation of the total bark proteins by DEAE-cellulose chromatography indicated that the final upsurge of bark proteins observed in November was associated primarily with one group of proteins (Peak III).     

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.