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Trees regenerating in the understory respond to increased availability of light caused by gap formation by undergoing a range of morphological and physiological adjustments. These adjustments include the production of thick, sun-type leaves containing thicker mesophyll and longer palisade cells than in shade-type leaves. We asked whether in the shade-regenerating tree Acer pseudoplatanus, the increase in leaf thickness and expansion of leaf tissues are possible also in leaves that had been fully formed prior to the increase in irradiance, a response reported so far only for a handful of species. We acclimated potted seedlings to eight levels (from 1 to 100%) of solar irradiance and, in late summer, transferred a subset of them to full sunlight. Within 30 days, the shaded leaves increased leaf mass per area and became thicker mostly due to elongation of palisade cells, except for the most shaded individuals which suffered irreversible photo-oxidative damage. This anatomical acclimation was accompanied by partial degradation of chlorophyll and a transient decline in photosynthetic efficiency of PSII (Fv/FM). These effects were related to the degree of pre-shading. The Fv/FM recovered substantially within the re-acclimation period. However, leaves of transferred plants were shed significantly earlier in the fall, indicating that the acclimation was not fully effective. These results show that A. pseudoplatanus is one of the few known species in which mature leaves may re-acclimate anatomically to increased irradiance. This may be a potentially important mechanism enhancing utilization of gaps created during the growing season.

The commonly observed negative relationship between stomatal density (SD) and atmospheric CO2 has led to SD being proposed as an indicator of atmospheric CO2 concentration. The use of SD as a proxy for CO2 , however, has been hampered by an insufficient understanding of the intraspecific variation of this trait. We hypothesized that SD in Pinus sylvestris, a widely distributed conifer, varies geographically and that this variation is determined by major climatic variables. By sampling needles from naturally growing trees along a latitudinal range of 32.25°, equivalent to 13.7°C gradient of mean annual temperature (MAT) across Europe, we found that SD decreased from the warmest southern sites to the coldest sites in the north at a rate of 4 stomata per mm2 for each 1°C, with MAT explaining 44% of the variation. Additionally, samples from a provenance trial exhibited a positive relationship between SD and the MAT of the original localities, suggesting that high SD is an adaptation to warm temperature. Our study revealed one of the strongest intraspecific relationships between SD and climate in any woody species, supporting the utility of SD as a temperature, rather than direct CO2 , proxy. In addition, our results predict the response of SD to climate warming.

Trees regenerating in the understory respond to increased availability of light caused by gap formation by undergoing a range of morphological and physiological adjustments. These adjustments include the production of thick, sun-type leaves containing thicker mesophyll and longer palisade cells than in shade-type leaves. We asked whether in the shade-regenerating tree Acer pseudoplatanus, the increase in leaf thickness and expansion of leaf tissues are possible also in leaves that are already fully formed, a response reported so far only for a handful of species. We acclimated potted seedlings to eight levels (from 1 to 100%) of solar irradiance and, in late summer, transferred a subset of them to full sunlight. Within 30 days, the pre-shaded leaves increased leaf mass per area and became thicker mostly due to the elongation of palisade cells, except for the most shaded individuals which suffered irreversible photo-oxidative damage. This anatomical acclimation was accompanied by a transient decline in photosynthetic efficiency of PSII (Fv/FM), the magnitude of which was related to the degree of pre-shading. The Fv/FM recovered substantially within the re-acclimation period. However, leaves of transferred plants were shed earlier in the fall, indicating that the acclimation was not fully effective. These results show that A. pseudoplatanus is one of the few known species in which mature leaves may re-acclimate anatomically to increased irradiance. This may be an important mechanism enhancing utilization of gaps created during the growing season.

Background and Aims The reliance on external support by lianas has been hypothesized to imply a reduction in the biomass cost of stem construction and root anchorage, and an increased investment in leaves, relative to self-supporting plants. These evolutionary trade-offs have not been adequately tested in an ontogenetic context and on the whole-plant scale. Moreover, the hypothesis may be extended to other potentially limiting resources, such as nitrogen (N.) Methods Plants belonging to five con-familiar pairs of temperate liana/shrub species were cultivated in 120 L barrels and sequentially harvested over up to three growing seasons. To account for the ontogenetic drift, organ biomass and nitrogen fractions were adjusted for plant biomass and N pool, respectively. Key Results Lianas invested, on average, relatively less biomass in the root fraction in comparison with shrubs. This was offset by only insignificant increases in leaf or stem investment. Even though liana stems and roots showed higher N concentration in comparison with shrubs, plant N distribution was mostly driven by, and largely matched, the pattern of biomass distribution. Lianas also showed a greater relative growth rate than shrubs. The differences between the growth forms became apparent only when ontogenetic drift was controlled for. These results were confirmed regardless of whether reproductive biomass was included in the analysis. Conclusions Our results suggest that temperate lianas, in spite of their diverse, species-specific resource distribution patterns, preferentially allocate resources to above-ground organs at the expense of roots. By identifying this trade-off and demonstrating the lack of a general trend for reduction in stem investment in lianas, we significantly modify the prevailing view of liana allocation strategies and evolutionary advantages. Such a resource distribution pattern, along with the cheap unit leaf area and stem unit length construction, situates lianas as a group close to the fast acquisition/rapid growth end of the life strategy spectrum.

In some lianas the use of trellis modifies within-plant biomass allocation and stimulates growth, however in other species trellis use may negatively affect growth indicating a cost associated with climbing. In order to improve access to light, lianas use other plants as climbing trellises. Whereas in shaded lianas climbing a trellis may enhance biomass gain by increasing leaf-level irradiance, we suspected that trellis use may also stimulate growth by triggering modifications in whole-plant allocation of biomass and nitrogen. We evaluated responses to trellis in three temperate lianas: Hedera helix, Celastrus orbiculatus and Wisteria floribunda. Lianas were grown outdoors in 120-L barrels and with ample space, with and without trellises. Biomass and nitrogen accumulation, distribution patterns and total stem length were determined after three and four seasons. Responses were adjusted for individual plant biomass. Liana responses to trellises were species-specific. In W. floribunda trellis use enhanced growth, consistently with the increased leaf mass fraction and leaf area ratio while biomass distribution to roots was reduced. Biomass distribution to stems and total stem length were increased by trellis use. In contrast, growth and biomass distribution in C. orbiculatus were not altered by trellis use. In H. helix growth was reduced in plants climbing on trellises in comparison with plants creeping on the ground, but biomass distribution pattern was not altered. Moreover, supported H. helix plants accumulated less nitrogen than did unsupported plants, suggesting a less efficient uptake of this nutrient. The ability to plastically modify growth and allocation patterns in response to trellis found in some lianas may constitute an important aspect of their ecological strategy. In other species, however, the switch to climbing may incur physiological costs resulting in lower growth.

Recent climatic changes have resulted in an increased frequency and prolonged periods of drought and strained water resources affecting plant production. We explored the possibility of reducing irrigation in a container nursery and studied the growth response of seedlings of economically important forest trees: broadleaf deciduous angiosperms Fagus sylvatica, Quercus petraea and evergreen conifers Abies alba and Pinus sylvestris. We also studied markers of water stress including modifications of biomass allocation, leaf anatomy, proline accumulation and expression of selected genes. Growth of the broadleaved deciduous species was more sensitive to the reduced water supply than that of conifers. Remarkably, growth of the shade tolerant Abies was not affected. Adjustment of biomass allocations was strongest in P. sylvestris, with a remarkable increase in allocation to roots. In response to water deficit both deciduous species accumulated proline in leaves and produced leaves with shorter palisade cells, reduced vascular tissues and smaller conduit diameters, but not conifers. Relative transcript abundance of a gene encoding a Zn-finger protein in Q. petraea and a gene encoding a pore calcium channel protein 1 in A. alba increased as water deficit increased. These findings suggest that in container nursery, the genetic selection can be initiated by water deficit. Our study shows major differences between functional groups in response to irrigation, with seedlings of evergreen conifers having higher tolerance than the deciduous species. This suggests that major water savings could be achieved by adjusting irrigation regime to functional group or species requirements.

Evergreen plants are an important component of many ecosystems of the world and occur in nu- merous evolutionary lineages. In this article we review phenotypic traits of evergreen woody angiosperms occurring in habitats that regularly experience frost. Leaf anatomical traits such as sclerenchymatic tissues or prominent cuticles ensure mechanical strength while often enhancing tolerance of water deficit. The low ratio of photosynthetic to nonphotosynthetic tissues as well as modified cell wall structure and nitrogen allocation patterns in evergreen leaves result in lower mass-based photosynthetic rate and photosynthetic nitrogen use efficiency in comparison with deciduous leaves. Their photosynthetic apparatus is adapted for the survival of frost in a down-regulated state with potential for photosynthetic activity in winter during periods of permissive temperatures. Leaf structure interacts with the mechanisms of frost survival. Stem xylem in evergreen plants tends to contain smaller diameter conduits incurring greater resistance to freeze/ thaw induced cavitation than in deciduous plants, although at the cost of reduced hydraulic efficiency. In contrast, no such differences in hydraulic conductivity have been documented at the leaf level. There is ev- idence for reduced structural plasticity of evergreen leaves in response to variability in irradiance, however photosynthetic downregulation occurs in mature leaves in response to self shading. Some evergreen species exhibit slow leaf development and "delayed greening", while in many species aging is also a very protracted process. Finally, evergreen leaves may participate in carbohydrate and, less obviously, in nitrogen storage for the support of spring shoot and foliage growth, although the importance of this function is under debate. In conclusion, the evergreen leaf habit is correlated with numerous structural and functional traits at the leaf and also at the stem level. These correlations may generate trade-offs that shape the ecological strategies of evergreen plants.

Background As proposed by Darwin, climbers have been assumed to allocate a smaller fraction of biomass to support organs in comparison with self-supporting plants. They have also been hypothesized to possess a set of traits associated with fast growth, resource uptake and high productivity. Scope In this review, these hypotheses are evaluated by assembling and synthesizing published and unpublished data sets from across the globe concerning resource allocation, growth rates and traits of leaves, stems and roots of climbers and self-supporting species. Conclusions The majority of studies offer little support for the smaller allocation of biomass to stems or greater relative growth rates in climbers; however, these results are based on small sized (

Spectra of leaf traits in northern temperate forest canopies reflect major differences in leaf longevity between evergreen conifers and deciduous broadleaf angiosperms, as well as plastic modifications caused by within-crown shading. We investigated (1) whether long-lived conifer leaves exhibit similar intra-canopy plasticity as short-lived broadleaves, and (2) whether global interspecific relationships between photosynthesis, nitrogen, and leaf structure identified for sun leaves adequately describe leaves differentiated in response to light gradients. We studied structural and photosynthetic properties of intra-tree sun and shade foliage in adult trees of seven conifer and four broadleaf angiosperm species in a common garden in Poland. Shade leaves exhibited lower leaf mass-per-area (LMA) than sun leaves; however, the relative difference was smaller in conifers than in broadleaves. In broadleaves, LMA was correlated with lamina thickness and tissue density, while in conifers, it was correlated with thickness but not density. In broadleaves, but not in conifers, reduction of lamina thickness was correlated with a thinner palisade layer. The more conservative adjustment of conifer leaves could result from a combination of phylogenetic constraints, contrasting leaf anatomies and shoot geometries, but also from functional requirements of long-lived foliage. Mass-based nitrogen concentration (Nmass) was similar between sun and shade leaves, and was lower in conifers than in deciduous broadleaved species. Given this, the smaller LMA in shade corresponded with a lower area-based N concentration (Narea). In evergreen conifers, LMA and Narea were less powerful predictors of area-based photosynthetic rate (Amax(area)) in comparison with deciduous broadleaved angiosperms. Multiple regression for sun and shade leaves showed that, in each group, Amax(mass) was related to Nmass but not to LMA, whereas LMA became a significant codeterminant of Amax(mass) in analysis combining both groups. Thus, a fundamental mass-based relationship between photosynthesis, nitrogen, and leaf structure reported previously also exists in a dataset combining within-crown and across-functional type variation. Electronic supplementary material The online version of this article (doi:10.1007/s00442-012-2279-y) contains supplementary material, which is available to authorized users.

Semidwarf barley cultivars have been created in the past decades featuring shortened straw length. The semidwarfness trait is desirable because plants with reduced height are more resistant to lodging and, as a consequence, have an improved harvest index. Accordingly, both breeding and molecular mapping experiments were undertaken to enhance and evaluate the performance of semidwarf barley lines. To study the morpho-anatomical effects of the denso locus twenty barley lines developed by single seed descent technique were analysed. We measured several macro and microscopic traits of aboveground organs. We detected reduction in leaf size, which at tillering stage was also reflected by smaller sizes of some, but not all, categories of epidermal cells. Lower cell division frequencies were deduced for cell files with unchanged cell lengths. Denso leaf blades were thinner and had smaller vascular bundles and narrower tracheal elements compared to the wild type. On the other hand, a lack of effect was noted e.g. for sizes of stomata and size of spike. Denso gene thus exerted a multilevel dwarfing effect on cells, tissues and organs although not all organs and cells were affected to the same extent, and cell size or division frequency were modified depending on cell type.

Seasonal variability of maximum quantum yield of PSII photochemistry (Fv/Fm) was studied in needles of Taxus baccata seedlings acclimated to full light (HL, 100% solar irradiance), medium light (ML, 18% irradiance) or low light (LL, 5% irradiance). In HL plants, Fv/Fm was below 0.8 (i.e. state of photoinhibition) throughout the whole experimental period from November to May, with the greatest decline in January and February (when Fv/Fm value reached 0.37). In ML seedlings, significant declines of Fv/Fm occurred in January (with the lowest level at 0.666), whereas the decline in LL seedlings (down to 0.750) was not significant. Full recovery of Fv/Fm in HL seedlings was delayed until the end of May, in contrast to ML and LL seedlings. Fv/Fm was significantly correlated with daily mean (Tmean), maximal (Tmax) and minimal (Tmin) temperature and Tmin was consistently the best predictor of Fv/Fm in HL and ML needles. Temperature averages obtained over 3 or 5 days prior to measurement were better predictors of Fv/Fm than 1- or 30-day averages. Thus our results indicate a strong light-dependent seasonal photoinhibition in needles of T. baccata as well as suggest a coupling of Fv/Fm to cumulative temperature from several preceding days. The dependence of sustained winter photoinhibition on light level to which the plants are acclimated was further demonstrated when plants from the three light environments were exposed to full daylight over single days in December, February and April and Fv/Fm was followed throughout the day to determine residual sensitivity of electron transport to ambient irradiance. In February, the treatment revealed a considerable midday increase in photoinhibition in ML plants, much less in HL (already downregulated) and none in LL plants. This suggested a greater capacity for photosynthetic utilization of electrons in LL plants and a readiness for rapid induction of photoinhibition in ML plants. Further differences between plants acclimated to contrasting light regimes were revealed during springtime de-acclimation, when short term regeneration dynamics of Fv/Fm and the relaxation of nonphotochemical quenching (NPQ) indicated a stronger persistent thermal mechanism for energy dissipation in HL plants. The ability of Taxus baccata to sustain winter photoinhibition from autumn until late spring can be beneficial for protection against an excessive light occurring together with frosts but may also restrict photosynthetic carbon gain by this shade-tolerant species when growing in well illuminated sites.

We tested the organogenetic capacity of floral buds of Mammillaria albicoma Boed. (Cactaceae). Buds were incubated on solid MS medium supplemented with 0.1 mg l−1 α-naphthaleneacetic acid (NAA) and 5.0 mg l−1 6-benzylaminopurine (BA). Callus growth was observed from the cut explant base and from within the perianth. These calli during subsequent subcultures to the same medium gave rise to adventitious shoots. Shoots formed also directly from the perianth, as confirmed by observations in the light microscope and scanning electron microscope (SEM). On transfer to a fresh medium, the shoots produced proliferating cultures. This is the first report of regeneration of cactus shoots from floral explants.

As part of fir reintroduction program in the Karkonosze Mountains (S. Poland) a study was undertaken to investigate growth and ecophysiological performance of young firs (Abies alba) under varying degrees of natural shade. Three-year-old fir seedlings were planted in five forest stands dominated by Picea abies, Fagus sylvatica, Betula pendula, Pinus sylvestris or Larix decidua. Over four growing seasons, annual apical increments became progressively greater under the light permeable canopy of Larix, but remained stunned under the deep shade of Picea, while intermediate irradiance at other sites resulted in intermediate growth responses. Similarly, apical dominance ratio was positively influenced by irradiance. These two contrasting sites were subsequently used for ecophysiological comparisons. Fir seedlings under the Larix canopy had consistently higher photosynthetic rates throughout the day compared to those under Picea, but in both stands light adapted quantum yield of PSII remained mostly above 0.8 indicating lack of photoinhibition. Quantum yield declined transiently only during sunflecks, which were more frequent under Larix. Light dependence of apparent electron transport rates in both sites was linear showing lack of light saturation, while slope differences between sites were only slight. In contrast, depression of PSII quantum yield by light was less pronounced under Larix, showing an acclimation to higher irradiance. Fir needles produced under Larix were thicker and contained more chlorophyll, carotenoid and nitrogen on area basis (but, except for nitrogen, not on dry mass basis) suggesting that structural modification is a major component of acclimation to light. On anatomical levels, needles produced under Larix had the traits typical for sun-acclimated ones, while those under Picea for shade-acclimated. The magnitude of modifications was, however, less than that reported in literature for sun and shade needles. Young seedlings were able to acclimate to higher irradiance under Larix without showing light stress symptoms. This suggests that success of stand regeneration may be enhanced by adjusting light penetration through the overstorey. Fir seedlings found best growth conditions under the relatively light permeable crowns of light-demanding tree species (Larix decidua, Pinus sylvestris) suggesting that stands dominated by these trees are more suitable for transformation using silver fir seedlings than those with shade-tolerant tree overstoreys.

In continuous light, the Crassulacean acid metabolism plant Kalanchoe daigremontiana Hamet et Perrier has a circadian rhythm of gas exchange with peaks occurring during the subjective night. The rhythm of gas exchange is coupled to a weak, reverse phased rhythm of quantum yield of photosystem II (Phi (PSII)). To test if the rhythm of Phi (PSII) persists in the absence of stomatal control, leaves were coated with a thin layer of translucent silicone grease which prevented CO2 and H2O exchange. In spite of this treatment, the rhythm of Phi (PSII) occurred with close to normal phase timing and with a much larger amplitude than in uncoated leaves. The mechanism underlying the Phi (PSII) rhythm in coated leaves can be explained by a circadian activity of phosphoenolpyruvate carboxylase (PEPC). At peaks of PEPC activity, the small amount of CO2 contained in the coated leaf could have become depleted, preventing the carboxylase activity of Rubisco and causing decreases in electron transport rates (observed as deep troughs of Phi (PSII) at 23-h in LL and at ca. 24-h intervals afterwards). Peaks of Phi (PSII) would be caused by a downregulation of PEPC leading to improved supply of CO2 to Rubisco. Substrate limitation of photochemistry at 23 h (trough of Phi (PSII)) was also suggested by the weak response of ETR in coated leaves to stepwise light enhancement. These results show that photosynthetic rhythmicity in K. daigremontiana is independent of stomatal regulation and may originate in the mesophyll.