Role of red carotenoids in photoprotection during winter acclimation in Buxus sempervirens leaves

The red leaf coloration of several plant species during autumn and winter is due to the synthesis of phenolic compounds such as anthocyanins or red carotenoids. The latter occur very rarely and are non-ubiquitous and taxonomically restricted compounds. The present study shows that the leaves of common box ( Buxus sempervirens L.) accumulate red carotenoids (eschscholtzxanthin, monoanhydroeschscholtzxanthin, anhydroeschscholtzxanthin) as a response to photoinhibitory conditions during winter acclimation. These compounds are produced in a coordinated manner with the operation of other photoprotective systems: accumulation and sustained deepoxidation of VAZ pigments with a concomitant decrease in maximal photochemical efficiency, accumulation of alpha-tocopherol and a gradual decrease on chlorophyll content. All these processes were reversed when the photosynthetic tissues were transferred from photoinhibitory winter conditions to room temperature for 9 days. Buxus leaves showed a large degree of phenotype variation in the degree of reddening, ranging from green to orange. The differences in colour pattern were mainly due to differences in the accumulation of red carotenoids and xanthophyll esters. Red pigments were mainly anhydroeschscholtzxanthin and esters of eschscholtzxanthin. Conversely to fruit or petal chromoplasts, the plastids of red leaves in this species are not the terminal differentiated state but are able to redifferentiate again to chloroplasts. Their photoprotective role during winter as a light screen system or as antioxidants, in a similar way to other red pigments, and their implications on the wide ecological tolerance of this evergreen species are discussed.     

The leaf anatomy of a broad-leaved evergreen allows an increase in leaf nitrogen content in winter

In temperate regions, evergreen species are exposed to large seasonal changes in air temperature and irradiance. They change photosynthetic characteristics of leaves responding to such environmental changes. Recent studies have suggested that photosynthetic acclimation is strongly constrained by leaf anatomy such as leaf thickness, mesophyll and chloroplast surface facing the intercellular space, and the chloroplast volume. We studied how these parameters of leaf anatomy are related with photosynthetic seasonal acclimation. We evaluated differential effects of winter and summer irradiance on leaf anatomy and photosynthesis. Using a broad-leaved evergreen Aucuba japonica , we performed a transfer experiment in which irradiance regimes were changed at the beginning of autumn and of spring. We found that a vacant space on mesophyll surface in summer enabled chloroplast volume to increase in winter. The leaf nitrogen and Rubisco content were higher in winter than in summer. They were correlated significantly with chloroplast volume and with chloroplast surface area facing the intercellular space. Thus, summer leaves were thicker than needed to accommodate mesophyll surface chloroplasts at this time of year but this allowed for increases in mesophyll surface chloroplasts in the winter. It appears that summer leaf anatomical characteristics help facilitate photosynthetic acclimation to winter conditions. Photosynthetic capacity and photosynthetic nitrogen use efficiency were lower in winter than in summer but it appears that these reductions were partially compensated by higher Rubisco contents and mesophyll surface chloroplast area in winter foliage.     

Contrasting patterns of photosynthetic acclimation and photoinhibition in two evergreen herbs from a winter deciduous forest

The relationship between the microclimate within an Oak-Hickory forest and photosynthetic characters of two resident evergreen herbs with contrasting leaf phenologies was investigated on a monthly basis for 1 full year. Heuchera americana has leaf flushes in the spring and fall, with average leaf life spans of 6–7 months. Hexastylis arifolia produces a single cohort of leaves each spring with a leaf life span of 12–13 months. We predicted that among evergreen plants inhabiting a seasonal habitat, a species for which the frequency of leaf turnover is greater than the frequency of seasonal extremes would have a greater annual range in photosynthetic capacity than a species that only produced a single flush of leaves during the year. Photosynthetic parameters, including apparent quantum yield, maximum photosynthetic capacity (P_max), temperature of maximum photosynthesis, photochemical efficiency of PSII and leaf nitrogen (N) and chlorophyll concentrations, were periodically measured under laboratory conditions in leaves sampled from natural populations of both species. Mature leaves of both species acclimated to changing understory conditions with the mean seasonal differences being significantly greater for Heuchera than for Hexastylis. Area based maximum photosynthetic rates at 25°C were approximately 250% and 100% greater in winter leaves than summer leaves for Heuchera and Hexastylis respectively. Nitrogen concentrations were highest in winter leaves. Chlorophyll concentrations were highest in summer leaves. Low P_max/N values for these species suggest preferential allocation of leaf nitrogen into non-photosynthetic pools and/or light-harvesting function at the expense of photosynthetic enzymes and electron transport components. Despite the increase in photosynthetic capacity, there was evidence of chronic winter photoinhibition in Hexastylis, but not in Heuchera. Among these ecologically similar species, there appears to be a trade-off between the frequency of leaf production and the balance of photosynthetic acclimation and photoinhibition.     

Photosynthesis of Conifers in Relation to Annual Growth Cycles and Dry Matter Production

Observations that deciduous and evergreen conifers growing in Britain show similar annual growth increments, despite marked differences in short-term growth rates, led to a comparative study of seasonal photosynthetic capacity in established trees of four coniferous species with contrasting growth habits. The photo-synthetic data were compared with seasonal changes in environmental parameters and chloroplast ultrastructure. The maximum net photosynthetic rates (at 20°C) recorded for Larix leptolepis were higher than those for the evergreen conifers when expressed on a leaf weight basis but not when expressed per unit leaf area. The photosynthetic efficiency of new needles in the evergreen species showed an overall decline from just after needle maturity until just before budbreak in their second season, after which photosynthetic rates recovered temporarily, approaching previous maximum levels. There was no obvious correlation between seasonal photosynthetic efficiency (at 20°C) on the one hand, and daily air, and (30 cm) ground temperatures on the other, and there was no obvious winter suppression of evergreen photosynthetic rates. Evergreen needles showed starch loss and some membrane changes with the onset of winter, but there was no evidence for wintertime chloroplast clumping or membrane disruption.     

Estimating contribution of anthocyanin pigments to osmotic adjustment during winter leaf reddening

The association between plant water stress and synthesis of red, anthocyanin pigments in leaves has led some plant biologists to propose an osmotic function of leaf reddening. According to this hypothesis, anthocyanins function as a solute in osmotic adjustment (OA), contributing to depression of osmotic potential (Ψ_π) and maintenance of turgor pressure during drought-stressed conditions. Here we calculate the percent contribution of anthocyanin to leaf Ψ_π during OA in two angiosperm evergreen species, Galax urceolata and Gaultheria procumbens. Both species exhibit dramatic leaf reddening under high light during winter, concomitant with declines in leaf water potential and accumulation of solutes. Data previously published by the authors on osmotic potential at full turgor (Ψ_π,100) of G. urceolata and G. procumbens leaves before and after leaf reddening were used to estimate OA. In vivo molar concentrations of anthocyanin, glucose, fructose, and sucrose measured from the same individuals were converted to pressure equivalents using the Ideal Gas Law, and percent contribution to OA was estimated. Estimated mean OA during winter was −0.7 MPa for G. urceolata and −0.8 MPa for G. procumbens. In vivo concentrations of anthocyanin (3–10 mM) were estimated to account for ∼2% of OA during winter, and comprised <0.7% of Ψ_π,100 in both species. Glucose, fructose, and sucrose combined accounted for roughly 50 and 80% of OA for G. urceolata and G. procumbens, respectively, and comprised ∼20% of Ψ_π,100. We observed that a co-occurring, acyanic species (Vinca minor) achieved similar OA without synthesizing anthocyanin. We conclude that anthocyanins represent a measurable, albeit meager, component of OA in red-leafed evergreen species during winter. However, due to their low concentrations, metabolic costliness relative to other osmolytes, and striking red color (unnecessary for an osmotic function), it is unlikely that they are synthesized solely for an osmoprotectant role.     

Annual photosynthetic activities of temperate evergreen and deciduous broadleaf tree species with simultaneous and successive leaf emergence in response to altitudinal air temperature

To explain why the composition of evergreen and deciduous forests changes along air temperature gradients, we measured several traits of single leaves from temperate deciduous and evergreen broadleaf trees with simultaneous and successive leaf emergence growing at different altitudes in the field. The parameters included seasonal net photosynthetic rate, longevity, mass per area, nitrogen content, and photosynthetic nitrogen-use efficiency. With decreasing altitude, the leaf longevity of deciduous broadleaf trees increased, whereas the maximum net photosynthetic rate decreased. In contrast, leaf longevity of evergreen broadleaf trees decreased, whereas the minimum net photosynthetic rate in winter increased. Along the air temperature gradient, the annual production of deciduous trees with simultaneous leaf emergence may be constant, because the integrated lifetime net photosynthetic rate (ILNPR) of a single leaf changed little. In comparison, deciduous trees with successive leaf emergence may show enhanced annual production with increasing air temperature, by increasing the total leaf number per branch and tree under an extended growing season. Temperate evergreen broadleaf tree species may also show increased annual production with increasing air temperature by sufficiently raising the number of the first-year leaves to the total leaves of branch and tree, which is accelerated by raising the integrated first-year net photosynthetic rate of the single leaf, despite little change in the ILNPR. With increasing air temperature from cool-temperate to warm-temperate zones, evergreen broadleaf tree species gain an advantage of the annual production over deciduous broadleaf tree species with simultaneous leaf emergence.     

Xanthophyll cycle pigment and antioxidant profiles of winter-red (anthocyanic) and winter-green (acyanic) angiosperm evergreen species

Leaves of many angiosperm evergreen species change colour from green to red during winter, corresponding with the synthesis of anthocyanin pigments. The ecophysiological function of winter colour change (if any), and why it occurs in some species and not others, are not yet understood. It was hypothesized that anthocyanins play a compensatory photoprotective role in species with limited capacity for energy dissipation. Seasonal xanthophyll pigment content, chlorophyll fluorescence, leaf nitrogen, and low molecular weight antioxidants (LMWA) of five winter-red and five winter-green angiosperm evergreen species were compared. Our results showed no difference in seasonal xanthophyll pigment content (V+A+Z g(-1) leaf dry mass) or LMWA between winter-red and winter-green species, indicating red-leafed species are not deficient in their capacity for non-photochemical energy dissipation via these mechanisms. Winter-red and winter-green species also did not differ in percentage leaf nitrogen, corroborating previous studies showing no difference in seasonal photosynthesis under saturating irradiance. Consistent with a photoprotective function of anthocyanin, winter-red species had significantly lower xanthophyll content per unit chlorophyll and less sustained photoinhibition than winter-green species (i.e. higher pre-dawn F(v)/F(m) and a lower proportion of de-epoxidized xanthophylls retained overnight). Red-leafed species also maintained a higher maximum quantum yield efficiency of PSII at midday (F'(v)/F'(m)) during winter, and showed characteristics of shade acclimation (positive correlation between anthocyanin and chlorophyll content, and negative correlation with chlorophyll a/b). These results suggest that the capacity for photon energy dissipation (photochemical and non-photochemical) is not limited in red-leafed species, and that anthocyanins more likely function as an alternative photoprotective strategy to increased VAZ/Chl during winter.     

Relationship between leaf reddening,ROS and antioxidants in <Emphasis Type="Italic">Buxus microphylla</Emphasis> during overwintering

To provide a theoretical basis for revealing the mechanism of winter leaf reddening in evergreen species, the relationships between winter leaf reddening, reactive oxygen species (ROS) and the antioxidant system of Buxus microphylla ‘Wintergreen’ were studied. The pigment changes, ROS production, lipid peroxidation and antioxidants activities of sun leaves during the reddening and regreening processes were investigated, using green shade leaves as controls. The carotenoids in the sun leaves increased linearly with reddening but decreased with the regreening. There was no significant difference in either the superoxide anions (O ₂ ^?· ) or malondialdehyde (MDA) changes between the sun and shade leaves, and their O ₂ ^?· contents were positively correlated with MDA. In contrast to the shade leaves, the sun leaves showed a trend in which the hydrogen peroxide (H₂O₂) changes were closely related to the reddening process and positively correlated with carotenoids content but not with the MDA content. A similar trend was observed for catalase (CAT) and ascorbate peroxidase (APX) activity between the sun and shade leaves, but superoxide dismutase (SOD) and peroxidase (POD) activity and the ascorbate (AsA) content differed between these two ecotypes. Furthermore, the sun leaves had higher CAT activity and AsA content than the corresponding shade leaves. These results suggested that H₂O₂ might play an important role in the winter reddening of sun leaves by promoting the accumulation of carotenoids. In addition, SOD, POD and AsA probably play a photoprotective role in winter-red sun leaves, while the changes in O ₂ ^?· , CAT and APX were independent of winter leaf reddening and were more likely responses to stress caused by low temperatures.     

Summer and winter sensitivity of leaves and xylem to minimum freezing temperatures: a comparison of co-occurring Mediterranean oaks that differ in leaf lifespan

Freezing sensitivity of leaves and xylem was examined in four co-occurring Mediterranean oaks (Quercus spp.) grown in a common garden to determine whether freezing responses of leaves and xylem were coordinated and could be predicted by leaf lifespan. Freezing-induced embolism and loss of photosynthetic function were measured after overnight exposure to a range of subzero temperatures in both summer and winter. Both measures were found to be dependent on minimum freezing temperature and were correlated with leaf lifespan and vessel diameter. The dependence of xylem embolism on minimum freezing temperature may result from the decline in water potential with ice temperature that influences the redistribution of water during freezing and leads to an increase in xylem tension. Winter acclimatization had a relatively small effect on the vulnerability to freezing-induced embolism, although leaf photosynthetic function showed a strong acclimatization response, particularly in the two evergreen species. Quercus ilex, the species with the longest leaf lifespan and narrowest vessel diameters, showed the highest freezing tolerance. This helps explain its ability to inhabit a broad range throughout the Mediterranean region. By contrast, the inability of the deciduous oaks to maintain photosynthetic and vascular function throughout the winter indicates a competitive disadvantage that may prevent them from expanding their ranges.     

Coordination of leaf N,anatomy, photosynthetic capacity,and hydraulics enhances evergreen expansive potential

Key message

Reduced leaf longevity, N-fixation, and enhanced hydraulic capacity combined support greater shifts in seasonal photosynthetic capacity of an expansive understory evergreen woody species relative to co-occurring less expansive evergreen species.

Abstract

Physiological functioning typically declines with increased leaf life span. While an evergreen leaf habit is generally associated with reduced leaf N, physiological capacity, and slower growth, most expansive woody species are evergreens and/or N fixers. An evergreen leaf habit enables year-round activity and less investment in carbon and nutrients, while N-fixation enhances photosynthetic capacity. Our objective was to compare anatomy and physiology of three woody evergreens Ilex opaca Aiton (Aquifoliaceae), Kalmia latifolia L. (Ericaceae), and Myrica cerifera (Myricaceae) of varying leaf longevity, N-fixation capability, and known expansive potential in a deciduous forest understory to determine if seasonal physiological performance integrated these factors. We hypothesized that I. opaca (non-expansive) and K. latifolia (moderately expansive), which have longer leaf longevities, would have reduced physiological performance compared to M. cerifera (expansive), which has shorter leaf longevity, and symbiotically fixes atmospheric N. Stomatal conductance to water vapor, photosynthetic and hydraulic capacities, specific leaf area, and leaf %N decreased with increasing leaf life span; however, trends among species were not consistent seasonally. While hydraulic capacity remained constant throughout the year, photosynthetic capacity did not. During the growing season, M. cerifera displayed photosynthetic capacity similar to deciduous species, yet, during the winter, photosynthetic capacity was similar to the slower-growing evergreens. Reduced leaf life span, enhanced hydraulic capacity, and nitrogen fixation support the seasonal shift in photosynthetic capacity observed in M. cerifera. This “hybrid” strategy enables M. cerifera to maximize productivity during months of optimal conditions, thereby promoting rapid growth and expansion in the understory.     

Optimal use of leaf nitrogen explains seasonal changes in leaf nitrogen content of an understorey evergreen shrub

Background and Aims

Understorey evergreen species commonly have a higher leaf nitrogen content in winter than in summer. Tested here is a hypothesis that such changes in leaf nitrogen content maximize nitrogen-use efficiency, defined as the daily carbon gain per unit nitrogen, under given temperature and irradiance levels.

Methods

The evergreen shrub Aucuba japonica growing naturally at three sites with different irradiance regimes in Japan was studied. Leaf photosynthetic characteristics, Rubisco and leaf nitrogen with measurements of temperature and irradiance monthly at each site were determined. Daily carbon gain was determined as a function of leaf nitrogen content to calculate the optimal leaf nitrogen content that maximized daily nitrogen-use efficiency.

Key Results

As is known, the optimal leaf nitrogen content increased with increasing irradiance. The optimal leaf nitrogen content also increased with decreasing temperature because the photosynthetic capacity per Rubisco decreased. Across sites and months, the optimal leaf nitrogen content was close to the actual leaf nitrogen content and explained the variation in actual leaf nitrogen by 64 %. Sensitivity analysis showed that the effect of temperature on optimal nitrogen content was similar in magnitude to that of irradiance.

Conclusions

Understorey evergreen species regulate leaf nitrogen content so as to maximize nitrogen-use efficiency in daily carbon gain under changing irradiance and temperature conditions.     

哀牢山和猫儿山中山常绿和落叶阔叶树光合特性对季节温度变化的响应

研究调查了云南哀牢山中山常绿阔叶林和广西猫儿山中山落叶水青冈林中共生的常绿和落叶阔叶树光合特性在2005年的季节变化,探讨了两地温度气候差异对植物优势度的影响。在夏季,同一生境下的常绿植物和落叶植物之间以及两个生境树种之间的光合速率没有显著差别。当温度下降时,两个生境内的常绿和落叶植物的光合速率都下降,这与气孔的部分关闭和光系统Ⅱ活性的下降有显著的关系。气孔的部分关闭有利于减少水分蒸腾损失,但落叶树气孔关闭程度明显高于常绿植物而且无法避免严重的光抑制。常绿植物通过气孔调节来提高光合水分利用效率和通过光系统Ⅱ活性调节来降低光合机构的损伤来维持叶片四季常绿,但是猫儿山中山森林冬季低温严重损害气孔和光系统Ⅱ的功能而使常绿植物生长受到严重制约并限制其在群落中成为优势种,与之相反,哀牢山中山森林冬季相对暖和导致常绿植物能积累更多的光合产物并促进其在群落中形成优势。     

Aboveground biomass allocation,leaf growth,and photosynthesis patterns in tundra plant forms in arctic Alaska

Summary Tundra plant growth forms can generally be characterized as consisting predominantly of low-growing perennial grasses and sedges, perennial herbaceous forbs, dwarf deciduous shrubs, and dwarf evergreen shrubs. Gross aboveground carbon allocation, leaf growth, and photosynthesis pattern studies were initiated to develop a quantitative understanding of the functional importance of these particular tundra growth forms. Photosynthetic capacities of 13 species were determined under standardized exposure conditions using a¹⁴CO₂ field system and ranged between 5 and 47 mg CO₂·g dry wt^-1·h^-1. These results, in conjunction with detailed leaf growth determinations, support the generalization that species with an evergreen growth form have lower photosynthetic capacities than species with a perennial graminoid, forb, or deciduous shrub growth form. However, these low photosynthetic capacities in evergreen shrubs are associated with relatively extended leaf longevities. Conversely, deciduous shrub forms exhibited high photosynthetic capacities, but were offset by relatively short leaf longevity periods. The perennial grasses, sedges, and forbs showed patterns intermediate to these. As a result, it appears that among tundra species of different growth form, photosynthetic capacity is inversely related to leaf longevity.     

Photosynthesis or persistence: nitrogen allocation in leaves of evergreen and deciduous Quercus species

Photosynthetic nitrogen use efficiency (PNUE, photosynthetic capacity per unit leaf nitrogen) is one of the most important factors for the interspecific variation in photosynthetic capacity. PNUE was analysed in two evergreen and two deciduous species of the genus Quercus. PNUE was lower in evergreen than in deciduous species, which was primarily ascribed to a smaller fraction of nitrogen allocated to the photosynthetic apparatus in evergreen species. Leaf nitrogen was further analysed into proteins in the water‐soluble, the detergent‐soluble, and the detergent‐insoluble fractions. It was assumed that the detergent‐insoluble protein represented the cell wall proteins. The fraction of nitrogen allocated to the detergent‐insoluble protein was greater in evergreen than in deciduous leaves. Thus the smaller allocation of nitrogen to the photosynthetic apparatus in evergreen species was associated with the greater allocation to cell walls. Across species, the fraction of nitrogen in detergent‐insoluble proteins was positively correlated with leaf mass per area, whereas that in the photosynthetic proteins was negatively correlated. There may be a trade‐off in nitrogen partitioning between components pertaining to productivity (photosynthetic proteins) and those pertaining to persistence (structural proteins). This trade‐off may result in the convergence of leaf traits, where species with a longer leaf life‐span have a greater leaf mass per area, lower photosynthetic capacity, and lower PNUE regardless of life form, phyllogeny, and biome.     

Seasonal variation in foliar carbon exchange in Pinus radiata and Populus deltoides: respiration acclimates fully to changes in temperature but photosynthesis does not

We investigated seasonal variation in dark respiration and photosynthesis by measuring gas exchange characteristics on Pinus radiata and Populus deltoides under field conditions each month for 1 year. The field site in the South Island of New Zealand is characterized by large day-to-day and seasonal changes in air temperature. The rate of foliar respiration at a base temperature of 10 °C ( R ₁₀) in both pine and poplar was found to be greater during autumn and winter and displayed a strong downward adjustment in warmer months. The sensitivity of instantaneous leaf respiration to a 10 °C increase in temperature ( Q ₁₀) was also greater during the winter period. The net effect of this strong acclimation was that the long-term temperature response of respiration was essentially flat over a wide range of ambient temperatures. Seasonal changes in photosynthesis were sensitive to temperature but largely independent of leaf nitrogen concentration or stomatal conductance. Over the range of day time growth temperatures (5–32 °C), we did not observe strong evidence of photosynthetic acclimation to temperature, and the long-term responses of photosynthetic parameters to ambient temperature were similar to previously published instantaneous responses. The ratio of foliar respiration to photosynthetic capacity ( R _d/ A _sat) was significantly greater in winter than in spring/summer. This indicates that there is little likelihood that respiration would be stimulated significantly in either of these species with moderate increases in temperature – in fact net carbon uptake was favoured at moderately higher temperatures. Model calculations demonstrate that failing to account for strong thermal acclimation of leaf respiration influences determinations of leaf carbon exchange significantly, especially for the evergreen conifer.     

Photosynthesis and Photoprotection in Overwintering Plants

Abstract: Seasonal differences in the capacity of photosynthetic electron transport, leaf pigment composition, xanthophyll cycle characteristics and chlorophyll fluorescence emission were investigated in two biennial mesophytes ( Malva neglecta and Verbascum thapsus ) that grow in full sunlight, and in leaves/needles of sun and shade populations of several broad-leafed evergreens and conifers (Vinca minor, Euonymus kiautschovicus, Mahonia repens, Pseudotsuga menziesii [Douglas fir], and Pinus ponderosa). Both mesophytic species maintained or upregulated photosynthetic capacity in the winter and exhibited no upregulation of photoprotection. In contrast, photosynthetic capacity was downregulated in sun leaves/needles of V. minor, Douglas fir, and Ponderosa pine, and even in shade needles of Douglas fir. Interestingly, photosynthetic capacity was upregulated during the winter in shade leaves/needles of V. minor, Ponderosa pine and Euonymus kiautschovicus. Nocturnal retention of zeaxanthin and antheraxanthin, and their sustained engagement in a state primed for energy dissipation, were observed largely in the leaves/needles of sun-exposed evergreen species during winter. Factors that may contribute to these differing responses to winter stress, including chloroplast redox state, the relative levels of source and sink activity at the whole plant level, and apoplastic versus symplastic phloem loading, are discussed.     

Relative growth rate of leaf biomass and leaf nitrogen content in several mediterranean woody species

In many plant species, herbivory is a major determinant of leaf mortality and it can cause a strong reduction in productive potential. Most predation occurs on young, expanding leaves. Thus, a rapid growth of the leaves can reduce the impact of predation. Furthermore, in cold Mediterranean climates the length of the growing season is constrained to a short period in spring and early summer owing both to low winter temperatures and drought stress in early summer. Therefore, a rapid deployment of leaf area and a high photosynthetic capacity during the spring and early summer might have important positive effects on the final carbon balance of the leaf population. Relative growth rates (RGR) of leaf biomass were measured in 19 woody species typical of Central Western Spain with deciduous and evergreen habits. Highly significant differences were detected in the leaf growth rate of the different species. The differences between species, however, did not correlate either with the mean leaf life-span of each of the species or with other leaf traits such as photosynthetic capacity, specific leaf area or nitrogen content. Leaf growth rate was positively correlated with time elapsed between leaf initiation and fruit maturation, so that species with fruit dispersal in spring and early summer in general had lower leaf growth rates than species with autumn fruit shedding. This relationship shows the effects of the concurrence between vegetative and reproductive organs for nutrients and other resources. Nitrogen concentration in the leaves was very high at the time of bud break, and declined during leaf expansion owing to the dilution associated with the increase in structural components. The rate of nitrogen dilution was, thus, positively related to the leaf growth rate. Relative growth rates calculated for nitrogen mass in leaves were very low compared to the growth in total mass. This suggests that most leaf nitrogen is translocated from the plant stores to the leaf biomass before the start of leaf expansion and that the contribution of root uptake during leaf expansion is comparatively low.     

Leaf litterfall patterns of perennial plant species in the arid Patagonian Monte,Argentina

The objective of this study was to investigate the variation in leaf litterfall patterns of desert plant species in relation to the intra- and interannual variation of precipitation. We collected the leaf litterfall of 12 representative species of the dominant life forms in the arid Patagonian Monte (evergreen shrubs, deciduous shrubs, and perennial grasses) at monthly intervals during three consecutive years. All shrub species showed a marked seasonality in the pattern of leaf litterfall, but the date of the peak of leaf litterfall differed among them. The peak of leaf litterfall in three deciduous and three evergreen shrubs occurred in summer months while in one deciduous shrub and in two other evergreen shrubs the peak of leaf litterfall was in autumn and winter, respectively. In contrast, the leaf litterfall of perennial grasses occurred through the year without a seasonal pattern. In most shrub species, increasing annual precipitation was related to increasing leaf litterfall and the peak of leaf litterfall was positively related to precipitation events occurred some months before, during winter. Moreover, the magnitude of responses in terms of variation in leaf litterfall in relation to interannual variation of precipitation was not the same for all species. Evergreen shrubs showed lower responses than deciduous species. These differences in leaf litterfall patterns were consistent with differences in leaf traits. In conclusion, we found new evidence of species-specific responses of leaf litterfall patterns to precipitation, suggesting that other factors than precipitation may control leaf litterfall in desert plants.     

Variation in evergreen and deciduous species leaf phenology in Assam, India

In the present study phenological activities such as leaf and shoot growth, leaf pool size and leaf fall were observed for 3 years (March 2007–March 2010) in 19 tree species (13 evergreen and 6 deciduous species) in a wet tropical forest in Assam, India. The study area receives total annual average rainfall of 2,318 mm of which most rain fall (>70 %) occurs during June–September. Both the plant groups varied significantly on most of the shoot and leaf phenology parameters. In general, growth in deciduous species initiated before the evergreen species and showed a rapid shoot growth, leaf recruitment and leaf expansion compared to evergreen species. Leaf recruitment period was significantly different between evergreen (4.2 months) and deciduous species (6.8 months). Shoot elongation rate was also significantly different for evergreen and deciduous species (0.09 vs. 0.14 cm day^?1 shoot^?1). Leaf number per shoot was greater for deciduous species than for evergreen species (34 vs. 16 leaves). The average leaf life span of evergreen species (328 ± 32 days) was significantly greater than that of deciduous species (205 ± 16 days). The leaf fall in deciduous species was concentrated during the winter season (Nov–Feb), whereas evergreens retained their leaves until the next growing season. Although the climate of the study area supports evergreen forests, the strategies of the deciduous species such as faster leaf recruitment rate, longer leaf recruitment time, faster shoot elongation rate during favorable growing season and short leaf life span perhaps allows them to coexist with evergreen species that have the liberty to photosynthesize round the year. Variations in phenological strategies perhaps help to reduce the competition among evergreen and deciduous species for resources in these forests and enable the coexistence of both the groups.     

Niche differentiation with respect to light utilization among coexisting dwarf shrubs in a subarctic woodland

Summary Canopy structure, shoot design, and photosynthetic light recruitment were used to compare four coexisting dwarf shrub species with respect to light utilization. All four species showed different shoot designs which probably result in different light interception properties. Leaves of Vaccinium uliginosum showed the highest levels of photosynthetic light saturation but in situ the shoots of this species reached their maximum photosynthetic rate at the lowest photon flux densities. No consistent differences with respect to photosynthetic light responses were found between deciduous and evergreen species. At sites dominated by one of the deciduous species (Vaccinium uliginosum or V. myrtillus), the two evergreen species studied (V. vitis-idaea and Empetrum hermaphroditum) occurred in the understory, i.e., with their leaf distribution slightly below that of the deciduous species. Sites dominated by one of the evergreen species showed less vertical differentiation in leaf distribution between species.