Needle Senescence Research Articles

Comprehensive Proteome and Acetylome Analysis of Needle Senescence in Larix gmelinii.

Leaf senescence is essential for the growth and development of deciduous trees in the next season. Larix gmelinii, a deciduous coniferous tree, exhibits its most distinctive feature by turning yellow in the autumn and eventually shedding its leaves, resulting in significant changes in its appearance during the fall. Lysine acetylation plays an important role in diverse cellular processes; however, limited knowledge is available regarding acetylations in the needle senescence of L. gmelinii. In this study, the proteomics and acetylated modification omics of two phenotypic leaves, yellow and green (senescent and non-senescent) needles, were analyzed before autumn defoliation. In total, 5022 proteins and 4469 unique acetylation sites in 2414 lysine acylated proteins were identified, and this resulted in the discovery of 1335 differentially expressed proteins (DEPs) and 605 differentially expressed acetylated proteins (DAPs) in yellow versus green needles. There are significant differences between the proteome and acetylome; only 269 proteins were found to be DEP and DAP, of which 136 proteins were consistently expressed in both the DEP and DAP, 91 proteins were upregulated, and 45 proteins were down-regulated. The DEPs participate in the metabolism of starch and sucrose, while the DAPs are involved in glycolysis and the tricarboxylic acid cycle. Among them, DEPs underwent significant changes in glycolysis and citric acid cycling. Most of the enzymes involved in glycolysis and the citrate cycle were acetylated. DAPs were down-regulated in glycolysis and up-regulated in the citrate cycle. In all, the results of this study reveal the important role of lysine acetylation in the senescence of L. gmelinii needles and provide a new perspective for understanding the molecular mechanism of leaf senescence and tree seasonal growth.

Remotely sensed and ground measurements reveal intraspecific differences in early season needle unfolding and senescence, but lack of variability in litter flammability of Pinus halepensis

Pinus halepensis Mill. is a conifer typical of Mediterranean pinewoods adapted to drought and fire. Although the timing of seasonal events in P. halepensis is strongly associated with these stressors and other environmental factors, needle phenology and litter flammability are insufficiently characterized at the intraspecific level. These traits can be informative of the existence of locally adapted populations across the species’ distribution range, and remote sensing approaches are promising tools to infer phenological changes in forest trees. In this study, we investigated intraspecific differentiation of P. halepensis related to needle phenology at the onset of the fire season (May-June) through vegetation indexes (VIs) obtained from unmanned aerial vehicles. We collected drone images using multispectral and RGB sensors for 56 adult populations of P. halepensis categorized into five ecotypes growing in two common gardens located in Spain under contrasting conditions (dry-continental versus wet-coastal). In the dry trial, we additionally monitored the temporal variation of RGB-derived indexes using two flights spaced over a one-month period. We also performed four consecutive ground measurements of needle pigments and fuel moisture content and obtained litter flammability traits for a subset of populations. Most in situ measurements and flammability traits did not show population differentiation. Regarding RGB-derived VIs, we did not detect obvious temporal patterns which differed among populations. However, we observed greener canopies in June than in May, which are indicative of the pace of current-year needle development. We also deduced an earlier phenology (i.e., earlier current-year needle unfolding) in the dry-continental trial. Ecotypic differentiation was found for some vegetation indexes related to needle unfolding (i.e., TCARI/OSAVI) and old needle senescence (i.e., PRSI). These differences were generally consistent across trials and time, and fundamentally indicated that sub-humid ecotypes typical of the eastern Mediterranean showed earlier needle unfolding and old needle senescence than semiarid ecotypes thriving in western continental Mediterranean areas. The intraspecific divergence observed in early season phenology is potentially related to the existence of contrasting life-history strategies present at the intraspecific level for the species. In particular, some semi-arid ecotypes, which are known to optimize the trade-off between carbon gain and water loss, exhibited an early old needle senescence (high likelihood of crown fire development) coupled with a late needle unfolding (conservative water use). These results indicate a possible trade-off between drought and fire resistance for the species, which suggest the importance of considering the intraspecific characteristics of Aleppo pine in forest management actions.

Needle Senescence Affects Fire Behavior in Aleppo Pine (Pinus halepensis Mill.) Stands: A Simulation Study

Research Highlights: Pre-programmed cell death in old Aleppo pine needles leads to low moisture contents in the forest canopy in July, the time when fire activity nears its peak in the Western Mediterranean Basin. Here, we show, for the first time, that such needle senescence may increase fire behavior and thus is a potential mechanism explaining why the bulk of the annual burned area in the region occurs in early summer. Background and Objectives: The brunt of the fire season in the Western Mediterranean Basin occurs at the beginning of July, when live fuel moisture content is near its maximum. Here, we test whether a potential explanation to this conundrum lies in Aleppo pine needle senescence, a result of pre-programmed cell death in 3-years-old needles, which typically occurs in the weeks preceding the peak in the burned area. Our objective was to simulate the effects of needle senescence on fire behavior. Materials and Methods: We simulated the effects of needle senescence on canopy moisture and structure. Fire behavior was simulated across different phenological scenarios and for two highly contrasting Aleppo pine stand structures, a forest, and a shrubland. Wildfire behavior simulations were done with BehavePlus6 across a wide range of wind speeds and of dead fine surface fuel moistures. Results: The transition from surface to passive crown fire occurred at lower wind speeds under simulated needle senescence in the forest and in the shrubland. Transitions to active crown fire only occurred in the shrubland under needle senescence. Maximum fire intensity and severity were always recorded in the needle senescence scenario. Conclusions: Aleppo pine needle senescence may enhance the probability of crown fire development at the onset of the fire season, and it could partly explain the concentration of fire activity in early July in the Western Mediterranean Basin.

Drought tolerance and acclimation in Pinus ponderosa seedlings: the influence of nitrogen form.

Drought is a limiting factor to forest regeneration and restoration, which is likely to increase in intensity and duration under future climates. Nitrogen (N) nutrition is related to drought-resistance mechanisms in trees. However, the influence of chemical N form (inorganic and organic N) on physiological traits related to drought resistance has been sparsely studied in conifer seedlings. We investigated the effect of N forms on morpho-physiological traits of Pinus ponderosa Dougl. ex Laws. seedlings and subsequent influences in drought tolerance and acclimation. One-year-old seedlings were fertilized during 10weeks at 9mMN with different N forms [either NH4+, NO3- or organic N (amino acids mixture)] in their second year of growth. After fertilization, we measured traits associated with intrinsic drought tolerance (shoot water relations, osmotic regulation, photosynthesis and cell membrane stability). Seedlings were then subjected to an 8-week drought period at varying drought intensities to evaluate plant acclimation mechanisms. We demonstrated that P. ponderosa seedlings could efficiently use amino acids as a primary N source, showing similar performance to those grown with inorganic N forms. Nitrogen form influenced mainly drought-acclimation mechanisms rather than intrinsic drought tolerance. Osmotic potential at saturation (Ψπsat) was marginally affected by N form, and a significant relationship between proline concentration in needles and Ψπsat was found. During acclimation, seedlings fertilized with organic N minimized needle senescence, retained more nutrients in the oldest needles, had maximum increments in proline concentration and hastened the development of water-use efficiency mechanisms compared with those fertilized with inorganic N sources. Our results suggest an improved physiological drought acclimation of organic N-fertilized seedlings.

Representing explicit budburst and senescence processes for evergreen conifers in global models

Global ecosystem models lack an explicit representation of budburst and senescence for evergreen conifers despite their primordial role in the carbon cycle. In this study we evaluated eight different budburst models, combining forcing, chilling and photoperiod, for their ability to describe spring budburst, and one model of needle senescence for temperate evergreen coniferous forests. The models’ parameters were optimized against field observations from a national forest monitoring network in France. The best fitting budburst model was determined according to a new metrics which accounts for both temporal and spatial variabilities of budburst events across sites. The best model could reproduce observed budburst dates both at the site scale (±5 days) and at regional scale (±12 days). We also showed that the budburst models parameterized at site scale lose some predictive capability when applied at coarser spatial resolution, e.g., in grid-based simulations. The selected budburst model was then coupled to a senescence function defined from needle survivorship observations in order to describe the full phenology cycle of coniferous forests. Implemented in the process-driven ecosystem model ORCHIDEE, this new conifer phenology module represented accurately the intra and inter-annual dynamics of leaf area index at both the local and regional scales when compared against MODIS remote sensing observations. A sensitivity analysis showed only a small impact of the new budburst model on the timing of the seasonal cycle of photosynthesis (GPP). Yet, due to the faster renewal of needles compared to the standard version of ORCHIDEE, we simulated an increase in the GPP by on average 15% over France, while the simulated needle turnover was doubled. Compared to 1970–2000, projections indicated an advancement of the budburst date of 10.3 ± 2.8 and 12.3 ± 4.1 days in average over the period 2060–2100 with the best forcing and chilling-forcing models respectively. Our study suggests that including an explicit simulation of needle budburst and senescence for evergreen conifers in global terrestrial ecosystem models may significantly impact future projections of carbon budgets.

Impacts of White Pine Needle Damage on seasonal litterfall dynamics and wood growth of eastern white pine (Pinus strobus) in northern New England

White Pine Needle Damage (WPND) is a complex of foliar fungal pathogens that have established as a chronic disease impacting eastern white pine (Pinus strobus L.) stands in the northeastern United States. With long-term ecological and economic impacts in mind, it is critical to quantify the negative effects of this disease on tree and forest health in order to make informed management decisions. We measured litterfall to determine the timing and magnitude of WPND-induced defoliation across four study sites in the northeastern US between 2014–2016. We measured N concentrations of needles cast throughout the 2014 growing season to estimate total litter N flux resulting from WPND. Additionally, to quantify growth declines we measured annual basal area increment (BAI) from six symptomatic study sites in the infected region. We found that WPND-induced defoliation in the months of June and July accounted for 47% of the total annual litterfall across the study sites, often exceeding normal needle senescence in October. Foliar %N in June and July was 0.78 and 0.84% respectively, significantly higher than October concentrations of 0.40%, suggesting incomplete resorption of N during the summer months. Untimely summer defoliations resulted in a mean estimated N loss of 0.92 g N m−2 yr−1, representing 63% of the total growing season N input from foliage. Growth of symptomatic trees at all sites was reduced following outbreaks of WPND initiating between 2007–2009. Severely infected trees reduced BAI 25–73% compared to pre-outbreak years. Our results show that WPND-induced defoliation significantly alters litterfall and N dynamics of affected stands, and suggest that subsequent N limitation in addition to reduced foliar area greatly reduces annual wood growth within infected stands.

Growing season ecophysiology ofTaxodium distichum(L.) Rich. (bald cypress) saplings in a restored wetland: a baseline for restoration practice

Wetland restoration is a common practice in the southeastern United States, where changing land use patterns have resulted in the widespread degradation of coastal ecosystems. Re-establishment of indigenous vegetation is a fundamental paradigm of restoration practice. In freshwater wetlands, this often includes planting flood-tolerant species such as Taxodium distichum (L.) Rich (bald cypress). While much is known regarding the ecology of this foundational species, research establishing the ecophysiological profile of mature bald cypress under stress-free, baseline conditions is limited. Therefore, a study was undertaken in 2015 to monitor the growing season ecophysiology of a bald cypress stand in a restored coastal wetland. Net photosynthesis was constant in the early to mid-growing season, declining steadily late in the growth season until the visible onset of needle senescence. Patterns in transpiration matched those observed for photosynthesis, whereas needle conductance and chlorophyll fluorescence appeared to track expected patterns associated with needle development. Trees at this site appeared healthy and were reproductively competent. However, long-term monitoring of the stand would help evaluate the ecological sustainability of this restoration project. We advocate for the inclusion of ecophysiological-based vegetation monitoring within restoration projects, enabling a sensitive and early indication of the trajectory of a restoration plan.

Intraseasonal carbon sequestration and allocation in larch trees growing on permafrost in Siberia after 13C labeling (two seasons of 2013-2014 observation).

This research is an attempt to study seasonal translocation patterns of photoassimilated carbon within trees of one of the high latitudes widespread deciduous conifer species Larix gmelinii (Rupr. Rupr). For this purpose, we applied whole-tree labeling by 13CO2, which is a powerful and effective tool for tracing newly developed assimilates translocation to tissues and organs of a tree. Experimental plot has been established in a mature 105-year-old larch stand located within the continuous permafrost area near Tura settlement (Central Siberia, 64°17'13″N, 100°11'55″E, 148m a.s.l.). Measurements of seasonal photosynthetic activity and foliage parameters (i.e., leaf length, area, biomass, etc.), and sampling were arranged from early growing season (June 8, 2013; May 14, 2014) until yellowing and senescence of needles (September 17, 2013; September 14, 2014). Labeling by 13C of the tree branch (June 2013, for 3 branch replicates in 3 different trees) and the whole tree was conducted at early (June 2014), middle (July 2014), and late (August 2013) phase of growing season (for different trees in 3 replicates each time) by three pulses [(CO2)max=3000-4000ppmv, 13CO2 (30% v/v)]. We found at least two different patterns of carbon translocation associated with larch CO2 assimilation depending on needle phenology. In early period of growing season (June), 13C appearing in newly developed needles is a result of remobilized storage material use for growth purposes. Then approximately at the end of June, growth processes is switching to storage processes lasting to the end of growing season.

Vulnerability of low temperature induced needle retention in balsam fir (Abies balsamea L.) to vapor pressure deficits

Low temperature (LT) exposure has been shown to delay postharvest needle abscission and senescence (NAS) in balsam fir and changes in vapor pressure deficit (VPD) are expected to alter these processes. Two and half year-old seedlings were exposed to a LT of 5°C for 15 days while the control (CT) group was maintained at 22°C. Seedlings were then exposed to four different VPD (0.22, 0.87, 1.3 and 1.86 kPa) and observed for relative water content (RWC), xylem pressure potential (XPP), membrane injury index (MII), stem capacitance (SC) and NAS. An interactive effect of LT and VPD was observed in RWC, XPP, MII and NAS. Low VPD (0.22 and 0.87 kPa) resulted in less negative XPP, lower electrolyte leakage, higher SC and ultimately higher NAS than those at high VPD. Maximum NAS was recorded at 0.22 kPa. At 1.86 kPa, LT had 5× lower RWC (13%), 3× more negative XPP (−1.1 MPa), 1.8× higher membrane damage and 35% lower NAS (47 days) than CT. The SC declined with an increase in VPD with no effect of LT. The XPP and RWC of LT seedlings showed a positive relationship with NAS with R2 values of 0.54 and 0.59, respectively. LT offered no benefits to NAS at high VPD environments.

Pathogenicity of Phytophthora pluvialis to Pinus radiata and its relation with red needle cast disease in New Zealand

Background Red needle cast, a new foliage disease of Pinus radiata in New Zealand is described. The disease has been variable in incidence and severity both regionally and in different years. The early symptoms of discrete olive coloured lesions, often with a narrow dark resinous mark or band, were first recognised in winter of 2008 in plantation forests on the eastern coast of the North Island. These lesions develop further to result in rapid needle senescence and premature defoliation. The disease has been termed red needle cast in New Zealand as affected trees have a reddish appearance prior to the casting of the needles. The subsequent four years of monitoring have confirmed that, depending on location, symptoms are first observed in late autumn through late winter. Newly developing spring and summer foliage is seldom affected. Isolation from needles using a Phytophthora-selective medium frequently yielded an unknown species of Phytophthora which was subsequently found to be identical to Phytophthora pluvialis, a species described from Oregon, USA in 2013 where it is not associated with disease. Infection appears to be limited to the needles with no recoveries of Phytophthora pluvialis having been made from the roots, stems or branches. Occasionally a second species of Phytophthora, P. kernoviae, was also recovered from needles with the same symptoms.

Characterization of phytohormonal and postharvest senescence responses of balsam fir (Abies balsamea (L.) Mill.) exposed to short-term low temperature

To fulfill the US Thanksgiving and Christmas tree markets, balsam fir (Abies balsamea (L.) Mill.) is generally harvested before the cold season, anecdotally leading to premature needle senescence. Accordingly, we tested the hypothesis that LT exposure before harvest induces specific hormonal changes and delays postharvest senescence and/or abscission in balsam fir. Two hundred and six seedlings exposed to two temperature treatments for 48 h, LT at 5 °C and controls at 22 °C were severed off roots and monitored for their postharvest needle senescence. Root and shoot (needles and buds) tissues were examined for major endogenous hormone metabolites. LT increased shoot ABA (2,007 ng g−1 DW) by 2.5× and decreased GA44 (9.84 ng g−1 DW) by 3.5× over those in roots. LT did not alter cytokinins, auxins or any root hormonal concentration. With auxins, only IAA, IAA-Asp, IAA-Leu and IAA-Glu were detected and the concentrations of IAA and IAA-Asp in shoots were lower than those found in roots. Among cytokinins, shoot c-ZR (58.95 ng g−1 DW) and t-ZR (4.17 ng g−1 DW) were 3× higher than those in roots. Apart from GA44, GA9 (136.76 ng g−1 DW) was abundant in shoots. The PBL and PNL were 46 and 1.2 %, irrespective of treatments. LT seedlings held needles 11 days longer than the controls (122 days). In balsam fir, short-term LT exposure augmented ABA and decreased GA44 levels in shoots and delayed postharvest needle senescence.

Needle longevity as a criterion of response to a climatic fluctuation (so called heat wave) in Scots pine populations at early phases of ontogeny

The paper presents results of an assessment of needle age classes in Scots pine partial populations in the period of three and five years after outplanting in a lowland site in relation to conditions created by climatic elements. The method is based on Skuhravý’s (1987) description when the state of needle age classes of the third whorl (from the apex) is evaluated just once. Five-year summary of meteorological data was assessed according to the criteria of determination of periods of heat waves and dry spells. Differences in numbers of living needle age classes in the groups of Scots pine populations in the period of three and five years after their outplanting in permanent lowland sites proved the highest total number of living needle classes in the group of local lowland populations. On the contrary, the lowest total number of living needle classes was found out in the group of mountain populations. Critical climatic periods are overlaps of heat waves and dry spells that illustrate the complexity of moisture and temperature conditions of lowland sites. Although the Scots pine is a xerophytic species, it responded to the cumulation of adverse factors by accelerated needle senescence. Its physiological response to heat and dry stress consists in needle shedding, i.e. reduction in the assimilating area. As mountain populations are not usually exposed to heat waves, the impact of this phenomenon on them was the highest. On the other hand, the local (lowland) populations carrying information on the patterns of local climate from the reproduction period do not undergo any marked reduction in needle age classes.  

Complex Environmental Factors Affecting the Decline of <i>Pinus densiflora</i> in the Seto Inland Sea Area of Western Japan

This study reconsidered the cause of forest decline of Japanese red pine (Pinus densiflora Sieb. et Zucc.) in the Seto Inland Sea area, western Japan. Although the decline in pine forests was attributed to pine-wood nematode infection, the stress of air pollution was also shown to have negative physiological effects on pine needles. Secondary pollutants related to NOx accumulate in the morning dew then liquid-phase free radicals (•OH) form by photochemical reactions on the needle surface. The stress of polluted dew induces faster needle senescence and defective needle gas exchange. The influence of atmospheric O3 and SO2 was not significant. Understory vegetation also had negative physiological effects on the overstory pine needles through competition of roots for water and nutrients. Abandonment of management accelerates growth of understory vegetation causing decline of P. densiflora plantations. Each of these stresses was shown to independently and significantly reduce needle photosynthesis, which consequently seemed to reduce pine resistance to pathogens. In many cases, air pollution and progressing succession have not been considered as important causes of P. densiflora decline. Careful on-site atmospheric chemical and eco-physiological measurements are therefore important for accurate evaluations of the conditions of pine growth and the causes of decline.

Photochemical smog effects in mixed conifer forests along a natural gradient of ozone and nitrogen deposition in the San Bernardino Mountains

Toxic effects of photochemical smog on ponderosa and Jeffrey pines in the San Bernardino Mountains were discovered in the 1950s. It was revealed that ozone is the main cause of foliar injury manifested as chlorotic mottle and premature needle senescence. Various morphological, physiological and biochemical alterations in the affected plants have been reported over a period of about 40 years of multidisciplinary research. Recently, the focus of research has shifted from studying the effects of ozone to multiple pollutant effects. Recent studies have indicated that the combination of ozone and nitrogen may alter biomass allocation in pines towards that of deciduous trees, accelerate litter accumulation, and increase carbon sequestration rates in heavily polluted forests. Further study of the effects of multiple pollutants, and their long-term consequences on the mixed conifer ecosystem, cannot be adequately done using the original San Bernardino Mountains Air Pollution Gradient network. To correct deficiencies in the design, the new site network is being configured for long-term studies on multiple air pollutant concentrations and deposition, physiological and biochemical changes in trees, growth and composition of over-story species, biogeochemical cycling including carbon cycling and sequestration, water quality, and biodiversity of forest ecosystems. Eleven sites have been re-established. A comparison of 1974 stand composition with data from 2000 stand composition indicate that significant changes in species composition have occurred at some sites with less change at other sites. Moist, high-pollution sites have experienced the greatest amount of forest change, while dryer low-pollution sites have experienced the least amount of stand change. In general, ponderosa pine had the lowest basal area increases and the highest mortality across the San Bernardino Mountains.

Carbohydrate accumulation in Siberian spruce needles of various ages

The content of mono- and oligosaccharides, hemicelluloses, and water-soluble polysaccharides was studied in Siberian spruce (Picea obovata Ledeb.) needles of various ages. Accumulation of carbohydrates during the growth period and the ratio of their individual fractions were determined by the proportions between the growth rate and assimilate export in developing needles. Throughout the growth period, the content of monosaccharides in needles of all ages was much higher than that of oligosaccharides. Unlike the period of active growth, the dormancy period was characterized by a wider diversity of oligosaccharides and their rather high content. Hemicelluloses that accumulated during the previous growth season and water-soluble polysaccharides of 2- and 3-year-old needles were involved into metabolism at the time of active growth of current-year needles. Analysis of carbohydrate accumulation in 1- to 9-year-old needles showed that the monosaccharide concentration was higher in needles of the first two years. The content of hemicelluloses A and B and water-soluble polysaccharides was higher in older needles. During the growth period, the assimilates of 2- and 3-year-old needles were partly used for the formation of young needles. The increase in relative content of polysaccharides during needle senescence reflects the involvement of photosynthetic products into the secondary pool of carbon metabolites.

Nutritional and Sexual Responses of Jack Pine to Ammonium Nitrate and Gibberellins

Abstract The nutritional and sexual responses of jack pine (Pinus banksiana Lamb.) seed orchard trees to NH4NO3 fertilization and GA4/7 foliar sprays were examined in seedling and grafted orchards at an Ontario location where soils were low N acidic sandy loams and in a Saskatchewan grafted orchard on high N calcareous sandy clay loams. High seed strobilus abortion and low pollen strobilus production and seed yields characterized the Saskatchewan trees. NH4NO3 fertilizer increased soil NH4+ and NO3- at both locations, current-year needle N and Mg in Ontario, Fe in Saskatchewan, and Ca at both locations, seed strobilus initiation and seed yields in Ontario; it decreased K in Saskatchewan. GA4/7 increased needle senescence and current-year needle N and P at both locations, K at Saskatchewan, seed and pollen strobilus initiation in Ontario, seed strobilus initiation and abortion in Saskatchewan, and seed yields per tree in Ontario; it decreased Ca at both locations, Mg at Saskatchewan, and diameter increments the year following treatments in Ontario. NH4NO3 and GA4/7 treatments did not interact for strobilus counts. Sexual expression varied with clones in Saskatchewan but not with families in Ontario. Clones interacted with GA4/7 but not NH4NO3 for seed strobilus initiation. For. Sci. 45(1):136-153.

Photosynthetic decline and pigment loss during autumn foliar senescence in western larch (Larix occidentalis).

We measured needle pigment content and photosynthetic rates of 1-year-old western larch (Larix occidentalis Nutt.) during autumn foliar senescence. Chlorophyll (Chl) and carotenoid (xanthophyll + b-carotene) contents of needles declined 11 and 17%, respectively, before CO(2) assimilation rate began to decline. Chlorophyll a/b ratio, Chl/carotenoid ratio, photochemical efficiency (F(v)/F(m)), and photochemical quenching did not begin to decline until late in senescence. Internal CO(2)/ambient CO(2) did not change during needle yellowing. In seedlings in warmed soil (average 3 degrees C above natural conditions), the decline in needle chlorophyll content was delayed by 10 days and the decline in CO(2) assimilation rate was delayed by 5 days, compared with seedlings in soil at ambient temperature. In seedlings exposed to an extended 16-h photoperiod, the decline in needle chlorophyll content was delayed by 32 days, and the decline in CO(2) assimilation rate was delayed by 21 days, compared with seedlings exposed to natural day lengths. In addition to delaying the onset of needle senescence, the treatments affected the sequence of events during senescence. Differences among treatment groups provide evidence that the onset of pigment loss and photosynthetic decline and the sequence of events during needle senescence are affected by soil temperature and day length.

Nutrient retranslocation in the foliage of Pinus sylvestris L. growing along a heavy metal pollution gradient.

Retranslocation of N, P, K, Ca, Mg, Mn, Fe, Zn, Cu and Ni within the foliage of Scots pine (Pinus sylvestris L.) was studied during autumnal needle senescence along a heavy-metal pollution gradient in western Finland. The stands were located at distances of 0.5 (Har 0.5), 4 (Har 4) and 8 km (Har 8) to the southeast of a copper-nickel smelter at Harjavalta. A background study stand was located at Hämeenkangas (Häm), 60 km northeast of the smelter. During needle senescence, the mobile nutrients N, P and K decreased by 67-88% and needle dry mass decreased by 31-43%. Retranslocation of P and K was less efficient in the stand nearest the smelter (Har 0.5) than in the other stands. During needle senescence at stand Har 0.5, heavy metal content either decreased slightly (Zn, Ni) or increased (Fe, Cu), whereas in the background study stand, retranslocation efficiency of Cu was 75%. Net throughfall (the amount of a nutrient in throughfall minus the amount in open precipitation) was higher in stand Har 0.5 than in stand Har 8 for all of the elements measured, except N and P which were intercepted by the canopy. In stand Har 8, deposition of S, Cu and Ni was intercepted by the canopy, and net throughfall of Fe and Zn was very low. In contrast, S and heavy metal contents of net throughfall in stand Har 0.5 were greater than the decrease in these elements in the needles indicating that accumulation on needle surfaces and subsequent wash-off accounted for a major part of the fluxes. Furthermore, the Cu content in net throughfall during the senescence period was higher than the original Cu content of the needles in July, providing additional evidence that much of the needle Cu content was comprised of dry deposits of Cu on the needle surface.

Early needle senescence and thinning of the crown structure of Picea abies as induced by chronic SO2pollution. I. Model deduction and analysis

AbstractRegarding time ranges of years, a rationale has been developed which is capable of explaining observed ‘spruce decline’ symptoms observed when spruce is exposed to air containing ambient levels of SO2. It integrates and interrelates (i) ecophysiological data (tree morphology, assimilate partitioning, canopy turnover, senescence physiology, stomatal conductance, canopy throughfall, sulphur metabolism, tonoplast symport), (ii) pedological data (soil leaching, cation recycling, litter decomposition, forest nutrition), and (iii) meteorological data (site elevation, length of the annual trunk growth period, SO2‐pollution). Furthermore, it can explain field observations at numerous sites of spruce decline in central Europe where SO2 is implicated as a factor of forest decline: (i) thinning of the canopy structure; (ii) early needle senescence; (iii) cation deficiency; (iv) low SO2 tolerance at sites with depleted soils in the mountains; (v) synergism of SO2pollution and acidic precipitation; (vi) recovery after liming, fertilization and after decreasing SO2 pollution; and (vii) higher SO2 tolerances of deciduous angiosperms. Different SO2tolerance strategies are identified that are employed by more SO2‐tolerant tree species. Ecophysiological SO2tolerance factors interact in a complex synergistic or antagonistic manner. It is concluded that chronic SO2 pollution at ambient concentrations predisposes mainly evergreen gymnosperms to suffer under synergistic environmental stresses (frost, drought, pathogens, etc.). Thinning of the crown structure is massive at extreme sites, where several stresses act simultaneously on the trees (depleted soils, high SO2 pollution, acidic rain, etc.). Mathematical formulations allow precise definitions of terms such as cooperativity, synergism, antagonism, vitality, predisposition, latency, etc. This universal rationale, which is applicable to all tree species, is exemplified here for Norway spruce (Picea abies [L.] Karst.). Integration of parameters yields an ordinary differential equation, which can be solved analytically. It predicts reversible dynamics of crown structures and gives an ecophysiological background to‘damage’.

Early needle senescence and thinning of the crown structure of Picea abies as induced by chronic SO2 pollution. II. Field data basis, model results and tolerance limits

AbstractField data on the sulphur and cation budget of growing Norway spruce canopies (Picea abies [L.] Karst.) are summarized. They are used to test a spruce decline model capable of quantifying effects of chronic SO2 pollution on spruce forests. At ambient SO2 concentrations, acute SO2 damage is rare, but exposure to polluted air produces reversible thinning of the canopy structure with a half‐time of a few years. Canopy thinning in the spruce decline model is highest (i) at elevated SO2 pollution, (ii) in the mountains, (iii) at unfertilized sites with poor K+, Mg2+ or Zn2+ supply, (iv) at low spruce litter decomposition rates, and (v) acidic, shallow soils at high annual precipitation rates in the field and vice versa. Model application using field data from Würzburg (moderate SO2 pollution, alkaline soils, no spruce decline) and from the Erzgebirge (extreme SO2 pollution, acidic soils in the mountains, massive spruce decline) predicts canopy thinning by 2–11% in Würzburg and by 45–70% in the Erzgebirge. The model also predicts different SO2‐tolerance limits for Norway spruce depending on the site elevation and on the nutritional status of the needles. If needle loss of more than 25% (damage class 2) is taken to indicate ‘real damage’ exceeding natural variances, then for optimum soil conditions SO2 tolerance limits range from (27.3 ± 7.4) μg m−3 to (62.6 ± 16.5) μg m−3. For shallow and acidic soils, SO2 tolerance limits range from (22.0 ± 5.5) μg m−3 to (37.4 ± 7.5) μ m−3. These tolerance limits, which are calculated on an ecophysiological data basis for Norway spruce are close to epidemiological SO2‐toIerance limits as recommended by the IUFRO, UN‐ECE and WHO. The observed statistical regression slope of the plot (damaged spruce trees vs. SO2‐pollution) in west Germany is confirmed by modelling (6% error). Model application to other forest trees allows deduction of the observed sequence of SO2‐sensitivity: Abies > Picea > Pinus > Fagus > Quercus. Thus, acute phytotoxicity of SO2 seems not to be involved in ‘forest decline’. Chronic SO2‐pollution induces massive canopy thinning of Abies alba and Picea abies only at unfavourable sites, where natural stress factors and secondary effects of SO2pollution act together to produce tree decline.