Contrasting seasonal overlaps between primary and secondary growth are linked to wood anatomy in Mediterranean sub-shrubs.

Whole-plant approaches allow quantification of the temporal overlap between primary and secondary growth. If the amount of time available to grow is short, there may be a high temporal overlap between shoot growth and wood formation. We hypothesise that such overlap depends on the duration of the growing season and relates to wood anatomy. We evaluated wood anatomy, shoot longitudinal and radial growth rates, fine root production and the concentrations of non-structural carbohydrates (NSC) in the wood of six sub-shrub species growing in sites with contrasting climatic conditions (Lepidium subulatum, Linum suffruticosum, Salvia lavandulifolia, Satureja montana, Ononis fruticosa, Echinospartum horridum). Sub-shrub species living in sites with a short growing season displayed a high overlap between aboveground primary and secondary growth and formed wide vessels, whereas species from the warmest and driest sites presented the reverse characteristics. The highest overlap was linked to a rapid shoot extension and thickening through the enhanced hydraulic conductivity provided by wide vessels. The reductions in NSC concentrations when growth peaked were low or moderate, indicating that sub-shrubs accumulate NSC in excess, as do trees. The temporal overlap among primary and secondary growth in woody plants may be connected to the duration and rates of shoot and wood growth, which in turn depend on the vessel lumen area.

Is leaf dry matter content a better predictor of soil fertility than specific leaf area?

BACKGROUND AND AIMS: Specific leaf area (SLA), a key element of the 'worldwide leaf economics spectrum', is the preferred 'soft' plant trait for assessing soil fertility. SLA is a function of leaf dry matter content (LDMC) and leaf thickness (LT). The first, LDMC, defines leaf construction costs and can be used instead of SLA. However, LT identifies shade at its lowest extreme and succulence at its highest, and is not related to soil fertility. Why then is SLA more frequently used as a predictor of soil fertility than LDMC? METHODS: SLA, LDMC and LT were measured and leaf density (LD) estimated for almost 2000 species, and the capacity of LD to predict LDMC was examined, as was the relative contribution of LDMC and LT to the expression of SLA. Subsequently, the relationships between SLA, LDMC and LT with respect to soil fertility and shade were described. KEY RESULTS: Although LD is strongly related to LDMC, and LDMC and LT each contribute equally to the expression of SLA, the exact relationships differ between ecological groupings. LDMC predicts leaf nitrogen content and soil fertility but, because LT primarily varies with light intensity, SLA increases in response to both increased shade and increased fertility. CONCLUSIONS: Gradients of soil fertility are frequently also gradients of biomass accumulation with reduced irradiance lower in the canopy. Therefore, SLA, which includes both fertility and shade components, may often discriminate better between communities or treatments than LDMC. However, LDMC should always be the preferred trait for assessing gradients of soil fertility uncoupled from shade. Nevertheless, because leaves multitask, individual leaf traits do not necessarily exhibit exact functional equivalence between species. In consequence, rather than using a single stand-alone predictor, multivariate analyses using several leaf traits is recommended.

Stomatal vs. genome size in angiosperms: the somatic tail wagging the genomic dog?

BACKGROUND AND AIMS: Genome size is a function, and the product, of cell volume. As such it is contingent on ecological circumstance. The nature of 'this ecological circumstance' is, however, hotly debated. Here, we investigate for angiosperms whether stomatal size may be this 'missing link': the primary determinant of genome size. Stomata are crucial for photosynthesis and their size affects functional efficiency. METHODS: Stomatal and leaf characteristics were measured for 1442 species from Argentina, Iran, Spain and the UK and, using PCA, some emergent ecological and taxonomic patterns identified. Subsequently, an assessment of the relationship between genome-size values obtained from the Plant DNA C-values database and measurements of stomatal size was carried out. KEY RESULTS: Stomatal size is an ecologically important attribute. It varies with life-history (woody species < herbaceous species < vernal geophytes) and contributes to ecologically and physiologically important axes of leaf specialization. Moreover, it is positively correlated with genome size across a wide range of major taxa. CONCLUSIONS: Stomatal size predicts genome size within angiosperms. Correlation is not, however, proof of causality and here our interpretation is hampered by unexpected deficiencies in the scientific literature. Firstly, there are discrepancies between our own observations and established ideas about the ecological significance of stomatal size; very large stomata, theoretically facilitating photosynthesis in deep shade, were, in this study (and in other studies), primarily associated with vernal geophytes of unshaded habitats. Secondly, the lower size limit at which stomata can function efficiently, and the ecological circumstances under which these minute stomata might occur, have not been satisfactorally resolved. Thus, our hypothesis, that the optimization of stomatal size for functional efficiency is a major ecological determinant of genome size, remains unproven.

Leaf exchange in a Mediterranean shrub: water, nutrient, non-structural carbohydrate and osmolyte dynamics.

Leaf exchange is an abrupt phenological event that drastically modifies the morphology and physiology of the aerial portion of the plant. We examined if water and osmolyte differences between old leaves and new organs trigger leaf exchange, and whether the differences are closely linked to the resource resorption process in senescing leaves. We monitored concentrations of osmolyte, water, non-structural carbohydrate, nitrogen and potassium in senescing leaves and in emerging new leaves and inflorescences of a Mediterranean leaf exchanger (Cistus laurifolius L.) growing in NE Spain. Old leaves rehydrated markedly during most of the senescence process, which co-occurred with the extension of new shoots, suggesting the lack of a clear-cut switch in water supply from old to new organs. The accumulation of osmolytes in the early stage of leaf senescence might account for this rehydration. Osmolyte dynamics in old leaves depended largely on the progression of resource resorption from senescing organs but were mostly unrelated to water content during late senescence. We conclude that dehydration of old leaves is not a prerequisite for the triggering of leaf exchange. The finding that most nutrients and carbohydrates accumulated in new organs before senescing leaves massively exported resources, and the absence of relevant differences between the dynamics of old leaves at the base of inflorescences and those at the base of vegetative shoots, indicate that the nutrient and carbohydrate demands of new organs do not trigger leaf exchange.

Non-structural carbohydrates and nitrogen dynamics in mediterranean sub-shrubs: an analysis of the functional role of overwintering leaves.

Previous studies have led to contrasting results about the role of overwintering leaves as storage sites, which is related to leaf longevity and life-form. The aim of this study was to evaluate the functional role of the leaves of four species of Mediterranean sub-shrubs, with different leaf phenology, as sources of nitrogen (N) and non-structural carbohydrates (NSC) for shoot growth. The seasonal dynamics of the concentrations and pools of N and NSC were assessed monthly in the leaves and woody organs of each species. Overwintering and spring leaves served as N and NSC sources for shoot growth in the evergreen species analyzed, providing up to 73 % and 324 % of the N demand for spring and autumn growth, respectively. Excess autumn N was stored in woody structures which contributed to the N and NSC requirements of spring growth. In the winter deciduous species, woody organs were the main N source for spring growth, while current photosynthesis from immature brachyblasts seemed to be the main carbon (C) source. Due to their short lifespan, overwintering and spring leaves did not show several translocation processes throughout their life time, their contribution to new growth being made during senescence. The successive exchange of leaf cohorts displayed by Mediterranean sub-shrubs might serve as a mechanism to recycle N and C between consecutive cohorts as plants perform the pheno-morphological changes needed to adapt their morphology to the seasonality of their environment.

Costs of reproduction as related to the timing of phenological phases in the dioecious Shrub Pistacia lentiscus L.

Females of woody dioecious species usually devote more resources to reproduction than males. This may lead to a decrease in female survival and growth. The costs of reproduction, however, can be lightened through a number of mechanisms, as for example avoiding the temporal coincidence of reproduction and vegetative growth. The aim of this study was to evaluate whether males and females of P. lentiscus differ in the timing of their vegetative growth, and to assess whether the sequencing of vegetative growth and reproduction reduces reproductive costs. We monitored phenology in males and females. We also compared male and female allocation of nutrients and biomass in the branch, and the developmental stability of the growing shoots. We did this both prior to and at the end of the fruiting period. Males and females showed similar vegetative and flowering phenologies. Males invested more biomass in flowering, but the sexes showed equal vegetative biomass and nutrient content prior to the fruiting period. In female branches, no trade-off was found between fruit load and current-year vegetative growth. In P. lentiscus, avoiding the overlap of flowering, vegetative growth and fruiting probably contributes to reduce the immediate costs of reproductive efforts, both in males and females.

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.

Environmental constraints on phenology and internal nutrient cycling in the Mediterranean winter-deciduous shrub Amelanchier ovalis Medicus.

The functional adjustments of winter-deciduous perennials to Mediterranean conditions have received little attention. The objectives of this study were: (i) to determine whether Amelanchier ovalis, a winter-deciduous shrub of Mediterranean and sub-Mediterranean regions, has nutritional and phenological traits in common with temperate zone deciduous trees and shrubs and (ii) to determine the constraints of Mediterranean environmental conditions on these traits. Over two years, phenology and nitrogen, and phosphorus concentrations were monitored monthly in the crown of A. ovalis. Leaf longevity, survival and nutrient resorption from senescing leaves were used to infer nutrient use efficiency and retention times of nutrients within the crown. In A. ovalis, bud burst was much earlier than in temperate deciduous trees and shrubs. Most vegetative and reproductive growth occurred in spring. Limited phenological development took place during the summer drought period. Unexpectedly, leaf shedding was very gradual, which might be related to water shortages in summer. Leaf longevity, nutrient resorption from senescing leaves, and maximum leaf nutrient concentrations indicated that nutrient retention times were short and nutrient use efficiency was low compared to that found in temperate deciduous plants and co-occurring Mediterranean evergreens. A. ovalis exhibited phenological development appropriate for a Mediterranean climate, although its limited ability to retain nutrients likely restricts the types of sites that it can occupy.

Seasonal branch nutrient dynamics in two Mediterranean woody shrubs with contrasted phenology.

BACKGROUND AND AIMS: Mediterranean woody plants have a wide variety of phenological strategies. Some authors have classified the Mediterranean phanaerophytes into two broad phenological categories: phenophase-overlappers (that overlap resource-demanding activities in a short period of the year) and phenophase-sequencers (that protract resource-demanding activities throughout the year). In this work the impact of both phenological strategies on leaf nutrient accumulation and retranslocation dynamics at the level of leaves and branches was evaluated. Phenophase-overlappers were expected to accumulate nutrients in leaves throughout most of the year and withdraw them efficiently in a short period. Phenophase-sequencers were expected to withdraw nutrients progressively throughout the year, without long accumulation periods. METHODS: To test this hypothesis, variations in phenology and leaf NPK in the crown of a phenophase-overlapper Cistus laurifolius and a phenophase-sequencer Bupleurum fruticosum were monitored monthly during 2 years. KEY RESULTS: Changes in nutrient concentration at the leaf level were not clearly related with the different phenologies. Nitrogen and phosphorous resorption efficiencies were lower in the phenophase-overlapper, and accumulation-retranslocation seasonality was similar in both species. Changes in the branch nutrient pool agreed with the hypothesis that the phenophase-overlapper accumulated nutrients from summer until the bud burst of the following spring, recovering a large nutrient pool during massive leaf shedding. The phenophase-sequencer did not accumulate nutrients from autumn until early spring, achieving lower nutrient recovery during spring leaf shedding. CONCLUSIONS: It is concluded that phenological demands influence branch nutrient cycling. This effect is easier to detect by assessing changes in the branch nutrient pool rather than changes in the leaf nutrient concentration.