Seasonal dynamics of non-structural carbohydrates in two species of mediterranean sub-shrubs with different leaf phenology

The seasonal dynamics of non-structural carbohydrates in the woody organs of two co-existing mediterranean sub-shrubs were analyzed. The two species show different leaf phenology during summer: Linum suffruticosum, maintains many of its green leaves, while Lepidium subulatum sheds most of its leaves. These different leaf phenologies are related to different strategies with regard to summer stress. The maintenance of leaves in Linum is related to its stress tolerance while Lepidium avoids stress by shedding its leaves. The main objectives were to: (1) determine the differences in the seasonal dynamics of non-structural carbohydrates among the main woody organs of both species; (2) verify if differences in the leaf phenology, and hence in the strategy with regard to summer drought, lead to different seasonal patterns of carbohydrate storage and use between the two species; (3) compare the seasonal dynamics of carbohydrates of the two studied sub-shrubs with those of mediterranean trees and shrubs previously reported in the literature. The concentration of soluble sugars (SS), starch and total non-structural carbohydrates (TNC) were assessed monthly, over 17 months, in the main roots, stems and the transition zone between root and shoot systems of both species. Starch storage capacity and SS, starch and TNC pools were calculated. The seasonal pattern of carbohydrate accumulation was similar among the woody organs analyzed, but it differed with those reported for mediterranean trees and shrubs. The two species showed different pools and seasonal patterns of non-structural carbohydrate concentrations in its woody organ, which corresponded to their different extent of leaf shedding. The stress-avoider Lepidium accumulated starch during spring shoot growth as a carbon store for summer respiration and had low pools of SS, whereas the stress-tolerant Linum increased SS during summer drought to maintain photosynthetic activity during summer and had low starch pools and storage capacity. However, irrespective of their different leaf shedding patterns, both species had a similar relative variation of their TNC concentration, which contrasts with previous results on deciduous and evergreen woody species.     

Seasonal variability of dry matter content and its relationship with shoot growth and nonstructural carbohydrates

This study assesses how different phases of shoot growth underlie seasonal change in leaf and stem dry matter content (LDMC and SDMC, respectively) of 12 woody Mediterranean species. The relationship between LDMC and nonstructural carbohydrate (NSC) concentrations is also explored and the seasonal vs interspecies variability of LDMC compared. LDMC, SDMC and shoot elongation rate (SER) were measured on a monthly basis for a minimum of 12 months. Bud growth rate (BGR) and NSC concentrations were also assessed in several of the study species. LDMC and SDMC decreased during shoot elongation in spring and increased in summer, showing a significant negative correlation with SER, but were unrelated to BGR. Half of the species analysed showed a positive relationship between LDMC and NSC. Seasonal fluctuations of LDMC within species were higher than interspecies differences, and species ranking was significantly affected by the month of sampling, except during winter months. Seasonal changes in LDMC and SDMC are mainly related to shoot elongation phenology, and NSC sink-source relationships between old and growing organs can explain this relationship in some species. Owing to the high seasonal variability in LDMC, it is recommended that samples for comparative purposes should be collected as close to the winter as possible.     

Spring versus autumn leaf colours: Evidence for different selective agents and evolution in various species and floras

Red colouration is common in young and old leaves of broadleaf woody species. Assuming that leaf colours are adaptive, we examined, by comparing the colouration in young versus old leaves, the possibility that different selection agents may have operated on spring versus autumn leaf colouration. We observed spring versus autumn colouration in three very different woody floras (Finland, Japan and Israel) in order to allow for a broad ecological and evolutionary spectrum. The null hypothesis was that if the same selective agents operated in spring and autumn, it is expected that when spring leaves are red, they should always be red in autumn, and when spring leaves are green, they should be green or yellow in autumn. We found that green spring leaves are almost exclusively associated with yellow leaf colour at senescence in autumn. Species with red autumn leaves almost always have at least some red colouration in their spring leaves. However, about half of the species with red spring leaves have yellow autumn leaves. Brown autumn leaves were not common in the species we studied. As about half of the species with red spring leaves have yellow autumn leaves but not vice versa, we conclude that there are many cases in which the selecting agents for spring versus autumn leaf colour were not the same.     

Overwintering leaves of a forest-floor fern, Dryopteris crassirhizoma (Dryopteridaceae): a small contribution to the resource storage and photosynthetic carbon gain

BACKGROUND AND AIMS: Dryopteris crassirhizoma is a semi-evergreen fern growing on the floor of deciduous forests. The present study aimed to clarify the photosynthetic and storage functions of overwintering leaves in this species. METHODS: A 2-year experiment with defoliation and shading of overwintering leaves was conducted. Photosynthetic light response was measured in early spring (for overwintering leaves) and summer (for current-year leaves). KEY RESULTS: No nitrogen limitation of growth was detected in plants subjected to defoliation. The number of leaves, their size, reproductive activity (production of sori) and total leaf mass were not affected by the treatment. The defoliation of overwintering leaves significantly reduced the bulk density of rhizomes and the root weight. The carbohydrates consumed by the rhizomes were assumed to be translocated for leaf production. Photosynthetic products of overwintering leaves were estimated to be small. CONCLUSION: Overwintering leaves served very little as nutrient-storage and photosynthetic organs. They partly functioned as a carbon-storage organ but by contrast to previous studies, their physiological contribution to growth was found to be modest, probably because this species has a large rhizome system. The small contribution of overwintering leaves during the short-term period of this study may be explained by the significant storage ability of rhizomes in this long-living species. Other ecological functions of overwintering leaves, such as suppression of neighbouring plants in spring, are suggested.     

Effects of leaf emergence on leaf lifespan are independent of life form and successional status

Abstract The longevity of a leaf is related to the benefit that the plant is able to derive from it. This benefit varies among seasons and as more leaves emerge, such that leaf lifespan can be limited by canopy position rather than physiological age. Using interval‐censored failure time analysis, we investigate leaf lifespan for 34 Mediterranean species in a previously published dataset involving species with different life forms and functional strategies. Failure time regression models were used to determine leaf lifespan, and to investigate how these effects varied among species. Median lifespan estimated for each species with two methods differed by less than 10% on average, but varied from 0.02–19.5% depending on the shape of the underlying failure time distribution. Within shoots, later‐emerging leaves had shorter lifespans for species with longer periods of leaf emergence, and the reverse was true for species with short emergence. Having accounted for the within‐shoot effect, leaves emerging in spring had shorter lifespans, particularly herbaceous species, whereas the reverse was true woody species. These effects were consistent among life forms and successional stages, and consistent with theories of within‐shoot translocation of resources following self‐shading.     

Relationships between phenology and the remobilization of nitrogen, phosphorus and potassium in branches of eight Mediterranean evergreens

Few studies have examined the effects of plant growth on nutrient remobilization in phenologically contrasting species. Here we evaluated the consequences of above-ground seasonality of growth and leaf shedding on the remobilization of nutrients from branches in eight evergreen Mediterranean phanaerophytes that differ widely in phenology. Vegetative growth, flower bud formation, flowering, fruiting, leaf shedding, and the variations in nitrogen (N), phosphorus (P) and potassium (K) pools in branches throughout the year were monitored in each species. Nitrogen and P remobilization occurred in summer, after vegetative growth and synchronously with leaf shedding. Despite the time-lag between growth and remobilization, the branches that invested more nutrients in vegetative growth also remobilized more nutrients from their old organs. Potassium remobilization peaked in the climatically harshest periods, and appears to be related to osmotic requirements. We conclude that N and P remobilization occurs mainly associated with leaf senescence, which might be triggered by factors such as the replenishment of nutrient reserves in woody organs, the hormonal relations between new and old leaves, or the constraints that summer drought poses on the amount of leaf area per branch in summer.     

Consequences of the Elimination of Old Leaves upon Spring Phenological Events and the New Leaves Nutrient Concentration in a Wintergreen Woody Species in the Southern Hemisphere

Previous studies analyzed the importance of old leaves conservancy for wintergreen species plant growth only after early spring old leaves elimination. However, carbon and nutrient resources for growth could have already been translocated from old leaves to shoots during autumn. In this work, the effect of old leaves absence on the leaf mass per area (LMA, g m⁻²) and nutrient concentration of new spring leaves, shoot growth, and flowering was studied in Aristotelia chilensis, an Andean Patagonic woody wintergreen species of Argentina. Plants were studied after autumn defoliation (AD) or late winter defoliation (WD) and results were compared to those of undamaged control plants (CO). The new leaves LMA and mineral nutrient (N, P, K, and Mg) concentration values did not decrease in AD or WD compared to CO plants. Conversely, CO plants showed higher flowering intensity and shoot lengthening compared to AD or WD plants. There were not remarkable differences regarding the defoliation time, though non-flowering shoots grew in a lesser degree than the flowering shoots in WD plants. It was concluded that A. chilensis old leaves cohort is an important source to shoot growth and flowering but their absence does not affect the new leaves structure or nutritional status from early spring in either AD or in WD plants. New leaves formation probably is guaranteed by resources (carbon and nutrients) previously stored in stems or even in the buds containing the preformed leaves since March, by the end of summer. Provided the availability of complete resources for the new leaf flush independently of the old leaves A. chilensis would restore the carbon balance as soon as possible to resume the growth of heterotrophic tissues at normal rates. Endogenous response to counterbalance the old leaves absence on non-flowering shoots was more effective when there was greater lag time between defoliation and shoot growth resume. Flowering and non-flowering shoots compete for the available resources when A. chilensis have not yet expanded leaves and shoots supporting reproductive structures were stronger sinks compared to non-flowering shoots in WD plants.     

Bud morphology and shoot growth dynamics in two species of Mediterranean sub-shrubs co-existing in gypsum outcrops

BACKGROUND AND AIMS: Understanding the effects of the environment on the morphology and shoot growth activities of plants is crucial to identifying plant ecological strategies. This study analysed the bud morphology, bud activity, shoot growth dynamics and shoot water content at full hydration (WC(h)) of two species of Mediterranean sub-shrubs, Lepidium subulatum and Linum suffruticosum, co-existing in gypsum outcrops in north-east Spain. METHODS: Sampling was conducted monthly over 2 years in one population per species. Buds were dissected under a stereo-microscope. Shoot growth was measured as the mean increase in shoot length of 15 marked individuals between two consecutive samplings. Bud activity was studied following the variations in the number of leaf primordia shorter than 1 mm and longer than 0.025 mm in the buds. KEY RESULTS: Both species bore naked buds and displayed discontinuous seasonal patterns of shoot growth, leaf primordia formation and WC(h). The number of leaf primordia in the bud peaked before the beginning of shoot expansion. In both species, organogenesis and expansion were uncoupled throughout the year. The time lapse between these two processes varied throughout the year, and was greatest for those elements differentiated in autumn. WC(h) was more closely related to shoot expansion than to organogenesis. CONCLUSIONS: Both species displayed similar bud morphology and similar seasonal patterns of bud and shoot growth, and WC(h) as a result of the strong seasonality of the Mediterranean climate in gypsum outcrops. The beginning of the spring period of expansion of long branches coincided with maximum values of WC(h), while the rest period of summer matched minimum values. These results support the hypothesis that the growth of long branches is strongly related to WC(h).     

Bud composition, branching patterns and leaf phenology in cerrado woody species

BACKGROUND AND AIMS: Plants have complex mechanisms of aerial biomass exposition, which depend on bud composition, the period of the year in which shoot extension occurs, branching pattern, foliage persistence, herbivory and environmental conditions. METHODS: The influence of water availability and temperature on shoot growth, the bud composition, the leaf phenology, and the relationship between partial leaf fall and branching were evaluated over 3 years in Cerrado woody species Bauhinia rufa (BR), Leandra lacunosa (LL) and Miconia albicans (MA). KEY RESULTS: Deciduous BR preformed organs in buds and leaves flush synchronously at the transition from the dry to the wet season. The expansion time of leaves is <1 month. Main shoots (first-order axis, A1 shoots) extended over 30 d and they did not branch. BR budding and foliage unfolds were brought about independently of inter-annual rainfall variations. By contrast, in LL and MA evergreen species, the shoot extension rate and the neoformation of aerial organs depended on rainfall. Leaf emergence was continuous for 2-6 months and lamina expansion took place over 1-4 months. The leaf life span was 5-20 months and the main A1 shoot extension happened over 122-177 d. Both evergreen species allocated biomass to shoots, leaves or flowers continuously during the year, branching in the middle of the wet season to form second-order (A2 shoots) and third-order (A3 shoots) axis in LL and A2 shoots in MA. Partial shed of A1 shoot leaves would facilitate a higher branching intensity A2 shoot production in LL than in MA. MA presented a longer leaf life span, produced a lower percentage of A2 shoots but had a higher meristem persistence on A1 and A2 shoots than LL. CONCLUSIONS: It was possible to identify different patterns of aerial growth in Cerrado woody species defined by shoot-linked traits such as branching pattern, bud composition, meristem persistence and leaf phenology. These related traits must be considered over and above leaf deciduousness for searching functional guilds in a Cerrado woody community. For the first time a relationship between bud composition, shoot growth and leaf production pattern is found in savanna woody plants.     

Seasonal patterns in species richness of herbivores: Macrolepidopteran larvae on Finnish deciduous trees

Abstract. 1. The seasonal distribution of macrolepidopteran species richness on Finnish deciduous trees vaned from positively skewed (peak in spring) to negatively skewed (peak in autumn).
2. The skewness values of species richness had a significant negative correlation ( r = - 0.98) with the duration of the seasonal shoot-growth period of the tree species.
3. Trees which complete their shoot growth early in the season ( Quercus type) produce new leaves only during spring, while trees whose shoot growth continues to autumn ( Populus type) do so throughout the summer.
4. Consequently, there is a difference in the number of available resources in the late summer foliage of different tree species, Trees ceasing leaf production early such as oak ( Quercus robur ) and bird cherry ( Prunus padus ) have one major resource type (mature leaves) in late-season foliage while trees like birches and alders have two Ooung and mature leaves).
5. Because young leaves formed late in the season are preferred to mature ones by some species of herbivores and because other species prefer mature leaves at the same time, the species richness of Populus-type trees is higher later in the season than the species richness of Quercus-type of trees, which have just one type of resource available.     

去叶对青杨雌雄植株生长和非结构性碳水化合物的影响

近年来,森林食叶害虫在全世界呈爆发趋势.树木的非结构性碳水化合物(NSC)如何响应叶片损失对其生长和生存至关重要.雌雄异株植物在维持森林生态系统稳定性方面扮演着重要角色.然而,目前对该类植物性别之间如何响应去叶的研究还比较少.本文以我国重要的经济和生态恢复树种青杨(Populus cathayana)为研究材料,比较了...     

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.     

Matter-economical roles of the evergreen foliage ofAucuba japonica,an understory shrub in the warm-temperate region of Japan

Leaf demography and productivity ofAucuba japonica, an understory shrub in the warm-temperate region, were examined and dry matter economy was analyzed to evaluate the roles of the evergreen foliage. Turnover of leaves occurred during a short period in spring. The mean leaf life span was about 2.6 years. Annual NAR (net assimilation rate) of each sample shoot was calculated from the biomass and the total dead mass estimated from scars of leaves and floral parts. The average NAR was 1.34±0.22 g·g⁻¹·yr⁻¹. The ratio of dry matter produced by leaves during their whole life span to the initial investment was 3.45±0.37. The annual NAR calculated for individual plants was negatively related to the life span of their leaves. The seasonal change in SLW (specific leaf weight) showed that the reserve material in leaves was accumulated from autumn to early spring and was consumed for the growth of new organs in the following season. The dry matter withdrawn in spring from the overwintering foliage amounted to 40% of dry mass of the new organs developed.     

Leaves of Field-Grown Mastic Trees Suffer Oxidative Stress at the Two Extremes of their Lifespan

Leaf senescence is a complex phenomenon occurring in all plant species, but it is still poorly understood in plants grown in Mediterranean field conditions and well-adapted to harsh climatic conditions. To better understand the physiological processes underlying leaf senescence in mastic trees (Pistacia lentiscus L.), we evaluated leaf growth, water and N content, photosystem II (PSII) photochemistry, lipid peroxidation and levels of photosynthetic pigments, antioxidants, abscisic acid, and salicylic acid and jasmonic acid during the complete leaf lifespan, from early expansion to late senescence in relation to natural climatic conditions in the field. While mature leaves suffered from water and N deficit during late spring and summer, both young (emerging) and old (senescing) leaves were most sensitive to photo-oxidative stress, as indicated by reductions in the F v/F m ratio and enhanced lipid peroxidation during late autumn and winter. Reductions in the F v/F m ratio were associated with low α-tocopherol (vitamin E) levels, while very old, senescing leaves additionally showed severe anthocyanin losses. We have concluded that both young (emerging) and old (senescing) leaves suffer oxidative stress in mastic trees, which may be linked in part to suboptimal temperatures during late autumn and winter as well as to low vitamin E levels.     

红色幼叶的适应意义探讨

很多木本植物的叶片会在春季和其他时期产生花青素而呈现红色, 该现象已经被众多学者所关注。本文对已有工作作了归纳总结。研究表明: 这种广泛存在的现象需要消耗营养和能量并影响光合作用, 并非只是代谢的副产品。前人以秋季红叶为研究对象, 主要提出了两大类假说: 第一类假说认为红叶是对强光、低温、干旱等恶劣环境的适应; 第二类认为红叶是植物通过化学防御警示、适口性差、隐蔽自身或暴露啃食者等方式来防御植食动物的啃食。这两类假说也都存在争议。目前对红色幼叶的研究相对较少且多侧重其作为独立视觉信号的作用, 而未能将红叶与植物的其他防御策略结合进行讨论。今后的研究应当综合环境因子的影响和啃食者的视觉分析, 并与植物其他出现红色的器官对比, 深入探讨红色幼叶的适应意义。     

圆柏属常绿木本植物元素含量的季节动态

测定分析了圆柏属2种常绿木本植物叶片干物质含量、叶绿素含量及N、P、K、Na、Ca、Mg等元素含量的季节变化和元素间的关系。结果发现,两种圆柏叶片N含量与叶绿素含量正相关,反映了其一年四季的生长情况;两树种N和K含量的季节变化趋势相似,夏、秋季高于冬、春季,表明两树种夏、秋季富集的营养物质较多,与夏、秋季较高的生物量一致;而P、Ca、Mg和Na含量是冬、春季较夏、秋季高,表明圆柏属植物通过积累这些无机渗透调节剂来增强其对低温的抗性。两树种叶片对N和K同步积累,对P、Na和Mg同步积累,但在元素积累模式上也存在一定的差异,元素间的关系也要复杂得多。     

秦岭东段栓皮栎叶片非结构性碳水化合物含量的季节动态

研究树木叶片非结构性碳水化合物(Nonstructural carbohydrate,NSC)组分的季节变化是掌握树木碳代谢规律的基础,也有利于判断以往研究仅凭生长季单次或几次(5次)的取样方法是否存在一定局限性。以秦岭东段栓皮栎(Quercus variabilis Blume)优势群落为研究对象,在其分布的海拔上下限(650 m和970 m),于2016年5月至2017年5月,通过月尺度周期性取样(共计9次),测定栓皮栎叶片NSC及其组分含量,并观测同期叶片物候变化。结果显示:(1)栓皮栎叶片NSC及其组分季节变化差异显著(P0.05),可溶性糖、淀粉和NSC变异系数分别为20.99%、52.28%和25.96%;(2)整体而言,栓皮栎叶片NSC最小值在展叶初期(3月末—4月初,5%左右),最大值在展叶末期(5月上旬,12%左右),之后NSC呈持续下降趋势。不同海拔NSC极值出现时间略有不同,叶片物候可能是影响年内极值的主要原因。(3)栓皮栎叶片NSC组成以可溶性糖为主(65%),这可能是树种在暖温带所采取的生长策略。(4)海拔对栓皮栎叶片NSC及其组分影响差异不显著,低海拔栓皮栎叶片NSC及其组分含量略大于高海拔。研究结果,栓皮栎叶片NSC含量存在明显的季节波动,适当加大NSC采样频率对于正确理解树木碳代谢十分必要。     

Seasonal patterns of leaf production in co‐occurring trees with contrasting leaf phenology: time and quantitative divergences

Neotropical savannas (‘cerrados’) of Central Brazil are characterized by the coexistence of a large diversity of tree species with divergent phenological behaviors, which reflect a great diversity in growth strategies. In the present study time behavior and quantitative aspects of shoot growth, shoot mortality, and leaf longevity and production were analyzed in 12 woody species of contrasting leaf phenology, adopting a functional group approach where 12 species were categorized into three functional groups: evergreen, decidous and brevideciduous, according to their leaf phenology. Shoot growth and leaf production were seasonal for the three functional groups, differing in their time of occurrence, but being concentrated during the last months of the dry season. Shoot growth differed between evergreens and deciduous, as well leaf production. Evergreens had higher rates of shoot growth, produced a higher number of leaves and had longer leaf longevity (around 500 days against 300 days in deciduous and brevideciduous). Leaf longevity was associated with patterns of leaf production when accounting for all phenological groups studied. It was possible to identify different patterns of aerial growth in savanna phenological groups, providing evidence of great functional variability amongst the groups studied.     

Biomass partitioning and leaf N,P - stoichiometry: comparisons between tree and herbaceous current-year shoots

We compare the biomass partitioning patterns and the nitrogen/phosphorus (N,P) stoichiometry of the current-year shoots of tree and herbaceous species and ask whether they scale in the same ways. Our analyses indicate that few statistically significant differences exist between the shoot biomass partitioning patterns of the two functional species-groups. In contrast, statistically significant N,P - stoichiometric differences exist between the two functional groups. Across all species, dry leaf mass scales nearly as the square of basal stem diameter and isometrically with respect to dry stem mass. However, total leaf N scales as the 1.37-power and as the 1.09-power of total leaf P across herbaceous and tree shoots, respectively. Therefore, tree shoots can be viewed as populations of herbs elevated by their older, woody herbaceous cohorts. However, tree leaf stoichiometry cannot be modelled in terms of herbaceous N,P - leaf stoichiometry.     

Shoot growth and seasonal changes of nonstructural carbohydrate concentrations at the upper and lower distribution limits of three conifers

Nonstructural carbohydrate (NSC) concentration in plant organs is an indicator of a balance between carbon sources (i.e., photosynthesis) and sinks (i.e., growth). Understanding how NSC concentrations change with altitude would help determine altitudinal changes in plant growth. This study compared shoot growth and seasonal changes in NSC concentrations of current-year and 1-year-old needles and branch woods between the upper and lower distribution limits of subalpine conifers Abies veitchii (1600–2000 m a.s.l.), A. mariesii (2000–2400 m a.s.l.), and Pinus pumila (2400–2800 m a.s.l.) in Japan. The lengths of 1-year-old shoots were shorter at the upper distribution limits for the three species, and concentrations and branch woods were all high in spring but decreased toward summer, increasing from summer to autumn. No clear difference was found for either parameter between upper and lower distribution limits for each species. Therefore, this study suggests that growth reduction at the upper distribution limits is due to reduction of both sink and source activities, with similar degrees for each species. However, further studies of sink and source activities, such as temperature-dependent photosynthesis and growth traits, are necessary to reveal clearly the cause of this growth reduction in high altitudes.