We determined the temporal dynamics of cambial activity and xylem cell differentiation of Scots pine (Pinus sylvestris L.) within a dry inner Alpine valley (750 m asl, Tyrol, Austria), where radial growth is strongly limited by drought in spring. Repeated micro-sampling of the developing tree ring of mature trees was carried out during 2 contrasting years at two study plots that differ in soil water availability (xeric and dry-mesic site).
In 2007, when air temperature at the beginning of the growing season in April exceeded the long-term mean by 6.4 °C, cambial cell division started in early April at both study plots. A delayed onset of cambial activity of c. 2 wk was found in 2008, when average climate conditions prevailed in spring, indicating that resumption of cambial cell division after winter dormancy is temperature-controlled. Cambial cell division consistently ended about the end of June/early July in both study years. Radial enlargement of tracheids started almost 3 wk earlier in 2007 compared with 2008 at both study plots. At the xeric site, the maximum rate of tracheid production in 2007 and 2008 was reached in early and mid-May, respectively, and c. 2 wk later, at the dry-mesic site. Since in both study years, more favorable growing conditions (i.e., an increase in soil water content) were recorded during summer, we suggest a strong sink competition for carbohydrates to mycorrhizal root and shoot growth. Wood formation stopped c. 4 wk earlier at the xeric compared with the dry-mesic site in both years, indicating a strong influence of drought stress on cell differentiation. This is supported by radial widths of earlywood cells, which were found to be significantly narrower at the xeric than at the dry-mesic site (P < 0.05).
Repeated cellular analyses during the two growing seasons revealed that, although spatial variability in the dynamics and duration of cell differentiation processes in Pinus sylvestris exposed to drought is strongly influenced by water availability, the onset of cambial activity and cell differentiation is controlled by temperature.
Cambium; dry inner Alpine valley; intra-annual growth; Scots pine; tracheid production; xylogenesis
•Background and Aims In the dry tropics, vegetative phenology varies widely with tree characteristics and soil conditions. The present work aims to document the phenological diversity of flowering and fruiting with reference to leafing events in Indian dry-tropical tree species.
•Methods Nine tree species, including one leaf-exchanging and eight deciduous showing varying leafless periods, were studied. Monthly counts of leaves, flowers and fruits were made on 160 tagged twigs on ten individuals of each species for initiation, completion and duration of different phenological events through two annual cycles.
•Key Results Variation in flowering relative to leaf flushing (which occurred just prior to or during a hot, dry summer) revealed five flowering types: summer flowering (on foliated shoots), rainy-season flowering (on foliated shoots following significant rains), autumn flowering (on shoots with mature leaves), winter flowering (on shoots undergoing leaf fall) and dry-season flowering (on leafless shoots). Duration of the fruiting phenophase was shortest (3–4 months) in dry-season and winter-flowering species, 6–9 months in rainy-and autumn-flowering species, and maximum (11 months) in summer-flowering species. A wide range of time lag (<1 to >8 months) between the start of vegetative (first-leaf flush) and reproductive (first-visible flower) phases was recorded in deciduous species; this time lag was correlated with the extent of the leafless period. A synthesis of available phenological information on 119 Indian tropical trees showed that summer-flowering species were most abundant (56 % of total species) amongst the five types recognized.
•Conclusions The wide diversity of seasonal flowering and fruiting with linkages to leaf flush time and leafless period reflect the fact that variable reproductive and survival strategies evolved in tree species under a monsoonic bioclimate. Flowering periodicity has evolved as an adaptation to an annual leafless period and the time required for the fruit to develop. The direct relationship between leafless period (inverse of growing period) and time lag between onset of vegetative and reproductive phases reflects the partitioning of resource use for supporting these phases. Predominance of summer flowering coupled with summer leaf flushing seems to be a unique adaptation in trees to survive under a strongly seasonal tropical climate.
Tropical tree phenology; flowering types; fruiting; asynchrony; leafless period; semi-evergreen species; summer flowering; summer leaf flushing
Background and Aims
Studies on xylogenesis focus essentially on the stem, whereas there is basically no information about the intra-annual growth of other parts of the tree. As roots strongly influence carbon allocation and tree development, knowledge of the dynamics of xylem production and maturation in roots at a short time scale is required for a better understanding of the phenomenon of tree growth. This study compared cambial activity and xylem formation in stem and roots in two conifers of the boreal forest in Canada.
Wood microcores were collected weekly in stem and roots of ten Abies balsamea and ten Picea mariana during the 2004–2006 growing seasons. Cross-sections were cut using a rotary microtome, stained with cresyl violet acetate and observed under visible and polarized light. The number of cells in the cambial zone and in differentiation, plus the number of mature cells, was counted along the developing xylem.
Xylem formation lasted from the end of May to the end of September, with no difference between stem and roots in 2004–2005. On the contrary, in 2006 a 1-week earlier beginning of cell differentiation was observed in the stem, with cell wall thickening and lignification in roots ending up to 22 d later than in the stem. Cell production in the stem was concentrated early in the season, in June, while most cell divisions in roots occurred 1 month later.
The intra-annual dynamics of growth observed in stem and roots could be related to the different amount of cells produced by the cambium and the patterns of air and soil temperature occurring in spring.
Abies balsamea; boreal forest; cambium; cell differentiation; cell wall thickening; lignification; Picea mariana; root; stem; xylem
We determined the temporal dynamic of cambial activity and xylem development of stone pine (Pinus cembra L.) throughout the treeline ecotone. Repeated micro-sampling of the developing tree ring was carried out during the growing seasons 2006 and 2007 at the timberline (1950 m a.s.l.), treeline (2110 m a.s.l.) and within the krummholz belt (2180 m a.s.l.) and the influence of climate variables on intra-annual wood formation was determined.
At the beginning of both growing seasons, highest numbers of cambial and enlarging cells were observed at the treeline. Soil temperatures at time of initiation of cambial activity were c. 1.5 °C higher at treeline (open canopy) compared to timberline (closed canopy), suggesting that a threshold root-zone temperature is involved in triggering onset of above ground stem growth.
The rate of xylem cell production determined in two weekly intervals during June through August 2006-2007 was significantly correlated with air temperature (temperature sums expressed as degree-days and mean daily maximum temperature) at the timberline only. Lack of significant relationships between tracheid production and temperature variables at the treeline and within the krummholz belt support past dendroclimatological studies that more extreme environmental conditions (e.g., wind exposure, frost desiccation, late frost) increasingly control tree growth above timberline.
Results of this study revealed that spatial and temporal (i.e. year-to-year) variability in timing and dynamic of wood formation of Pinus cembra is strongly influenced by local site factors within the treeline ecotone and the dynamics of seasonal temperature variation, respectively.
Cambium; intra-annual growth; Pinus cembra; temperature; tracheid production
Background and Aims
Our knowledge about the influences of environmental factors on tree growth is principally based on the study of dominant trees. However, tree social status may influence intra-annual dynamics of growth, leading to differential responses to environmental conditions. The aim was to determine whether within-stand differences in stem diameters of trees belonging to different crown classes resulted from variations in the length of the growing period or in the rate of cell production.
Cambial activity was monitored weekly in 2006 for three crown classes in a 40-year-old silver-fir (Abies alba) plantation near Nancy (France). Timings, duration and rate of tracheid production were assessed from anatomical observations of the developing xylem.
Cambial activity started earlier, stopped later and lasted longer in dominant trees than in intermediate and suppressed ones. The onset of cambial activity was estimated to have taken 3 weeks to spread to 90 % of the trees in the stand, while the cessation needed 6 weeks. Cambial activity was more intense in dominant trees than in intermediate and suppressed ones. It was estimated that about 75 % of tree-ring width variability was attributable to the rate of cell production and only 25 % to its duration. Moreover, growth duration was correlated to tree height, while growth rate was better correlated to crown area.
These results show that, in a closed conifer forest, stem diameter variations resulted principally from differences in the rate of xylem cell production rather than in its duration. Tree size interacts with environmental factors to control the timings, duration and rate of cambial activity through functional processes involving source–sink relationships principally, but also hormonal controls.
Cambial activity; forest-stand structure; silver fir (Abies alba); tree-ring formation; tree-to-tree competition; social status; wood anatomy; xylem cell differentiation
Within the alpine treeline ecotone tree growth is increasingly restricted by extreme climate conditions. Although intra-annual stem growth recorded by dendrometers can be linked to climate, stem diameter increments in slow-growing subalpine trees are masked by changes in tree water status.We tested the hypothesis that intra-annual radial stem growth in Pinus cembra is influenced by different climate variables along the treeline ecotone in the Austrian Alps. Dendrometer traces were compared with dynamics of xylem cell development to date onset of cambial activity and radial stem growth in spring.Daily fluctuations in stem radius reflected changes in tree water status throughout the treeline ecotone. Extracted daily radial increments were significantly correlated with air temperature at the timberline and treeline only, where budburst, cambial activity and enlargement of first tracheids also occurred quite similarly. A close relationship was detected between radial increment and number of enlarging tracheids throughout the treeline ecotone.We conclude that (i) the relationship between climate and radial stem growth within the treeline ecotone is dependent on a close coupling to atmospheric climate conditions and (ii) initiation of cambial activity and radial growth in spring can be distinguished from stem re-hydration by histological analysis.
dendrometer; Pinus cembra; radial increment; treeline ecotone; xylem formation
Background and Aims
Secondary growth via successive cambia has been intriguing researchers for decades. Insight into the mechanism of growth layer formation is, however, limited to the cellular level. The present study aims to clarify secondary growth via successive cambia in the mangrove species Avicennia marina on a macroscopic level, addressing the formation of the growth layer network as a whole. In addition, previously suggested effects of salinity on growth layer formation were reconsidered.
A 1-year cambial marking experiment was performed on 80 trees from eight sites in two mangrove forests in Kenya. Environmental (soil water salinity and nutrients, soil texture, inundation frequency) and tree characteristics (diameter, height, leaf area index) were recorded for each site. Both groups of variables were analysed in relation to annual number of growth layers, annual radial increment and average growth layer width of stem discs.
Between trees of the same site, the number of growth layers formed during the 1-year study period varied from only part of a growth layer up to four growth layers, and was highly correlated to the corresponding radial increment (0–5 mm year–1), even along the different sides of asymmetric stem discs. The radial increment was unrelated to salinity, but the growth layer width decreased with increasing salinity and decreasing tree height.
A patchy growth mechanism was proposed, with an optimal growth at distinct moments in time at different positions around the stem circumference. This strategy creates the opportunity to form several growth layers simultaneously, as observed in 14 % of the studied trees, which may optimize tree growth under favourable conditions. Strong evidence was provided for a mainly endogenous trigger controlling cambium differentiation, with an additional influence of current environmental conditions in a trade-off between hydraulic efficiency and mechanical stability.
Avicenia marina; cambial marking; mangrove; phloem; salinity; secondary growth; successive cambia; xylem
Seasonally dry tropical forests (SDTF) are characterized by pronounced seasonality in rainfall, and as a result trees in these forests must endure seasonal variation in soil water availability. Furthermore, SDTF on the northern Yucatan Peninsula, Mexico, have a legacy of disturbances, thereby creating a patchy mosaic of different seral stages undergoing secondary succession. We examined the water status of six canopy tree species, representing contrasting leaf phenology (evergreen vs. drought-deciduous) at three seral stages along a fire chronosequence in order to better understand strategies that trees use to overcome seasonal water limitations. The early-seral forest was characterized by high soil water evaporation and low soil moisture, and consequently early-seral trees exhibited lower midday bulk leaf water potentials (ΨL) relative to late-seral trees (−1.01 ± 0.14 and −0.54 ± 0.07 MPa, respectively). Although ΨL did not differ between evergreen and drought-deciduous trees, results from stable isotope analyses indicated different strategies to overcome seasonal water limitations. Differences were especially pronounced in the early-seral stage where evergreen trees had significantly lower xylem water δ18O values relative to drought-deciduous trees (−2.6 ± 0.5 and 0.3 ± 0.6‰, respectively), indicating evergreen species used deeper sources of water. In contrast, drought-deciduous trees showed greater enrichment of foliar 18O (∆18Ol) and 13C, suggesting lower stomatal conductance and greater water-use efficiency. Thus, the rapid development of deep roots appears to be an important strategy enabling evergreen species to overcome seasonal water limitation, whereas, in addition to losing a portion of their leaves, drought-deciduous trees minimize water loss from remaining leaves during the dry season.
Groundwater; Resource partitioning; Stable isotopes; Water-use efficiency; Yucatan Peninsula
Background and Aims
The timing of cambial reactivation plays an important role in the control of both the quantity and the quality of wood. The effect of localized heating on cambial reactivation in the main stem of a deciduous hardwood hybrid poplar (Populus sieboldii × P. grandidentata) was investigated.
Electric heating tape (20–22 °C) was wrapped at one side of the main stem of cloned hybrid poplar trees at breast height in winter. Small blocks were collected from both heated and non-heated control portions of the stem for sequential observations of cambial activity and for studies of the localization of storage starch around the cambium from dormancy to reactivation by light microscopy.
Cell division in phloem began earlier than cambial reactivation in locally heated portions of stems. Moreover, the cambial reactivation induced by localized heating occurred earlier than natural cambial reactivation. In heated stems, well-developed secondary xylem was produced that had almost the same structure as the natural xylem. When cambial reactivation was induced by heating, the buds of trees had not yet burst, indicating that there was no close temporal relationship between bud burst and cambial reactivation. In heated stems, the amount of storage starch decreased near the cambium upon reactivation of the cambium. After cambial reactivation, storage starch disappeared completely. Storage starch appeared again, near the cambium, during xylem differentiation in heated stems.
The results suggest that, in deciduous diffuse-porous hardwood poplar growing in a temperate zone, the temperature in the stem is a limiting factor for reactivation of phloem and cambium. An increase in temperature might induce the conversion of storage starch to sucrose for the activation of cambial cell division and secondary xylem. Localized heating in poplar stems provides a useful experimental system for studies of cambial biology.
Populus sieboldii × Populus grandidentata; localized heating, cambial reactivation; model system; storage starch; xylem differentiation
Background and Aims
The light availability on a temperate, deciduous-forest floor varies greatly, reflecting the seasonal leaf dynamics of the canopy trees. The growth and/or reproductive activity of understorey plants should be influenced by the length of the high-irradiance period from snowmelt to canopy closure. The aim of the present study was to clarify how spring-blooming species regulate the translocation of photosynthetic products to current reproduction and storage organs during a growing season in accordance with the changing light conditions.
Growth pattern, net photosynthetic rate, seed production, and shoot and flower production in the next year of Trillium apetalon were compared between natural and experimentally shaded conditions. Furthermore, translocation of current photosynthetic products within plants was assessed by a labelled carbon-chase experiment.
During the high-irradiance period, plants showed high photosynthetic ability, in which current products were initially used for shoot growth, then reserved in the rhizome. Carbon translocation to developing fruit occurred after canopy closure, but this was very small due to low photosynthetic rates under the darker conditions. The shading treatment in the early season advanced the time of carbon translocation to fruit, but reduced seed production in the current year and flower production of the next year.
Carbon translocation to the storage organ had priority over seed production under high-irradiance conditions. A shortened bright period due to early canopy closure effectively restricts carbon assimilation, which greatly reduces subsequent reproductive output owing to low photosynthetic products for fruit development and small carbon storage for future reproduction. As populations of this species are maintained by seedling recruitment, acceleration of canopy closure timing may influence the maintenance and dynamics of populations.
13C labelling; canopy closure; carbon translocation; deciduous forest; light availability; photosynthesis; spring-bloomer; Trillium apetalon
Flood-tolerant tree species of the Amazonian floodplain forests are subjected to an annual dry period of variable severity imposed when low river-water levels coincide with minimal precipitation. Although the responses of these species to flooding have been examined extensively, their responses to drought, in terms of phenology, growth and physiology, have been neglected hitherto, although some information is found in publications that focus on flooding.
The present review examines the dry phase of the annual flooding cycle. It consolidates existing knowledge regarding responses to drought among adult trees and seedlings of many Amazonian floodplain species.
Flood-tolerant species display variable physiological responses to dry periods and drought that indicate desiccation avoidance, such as reduced photosynthetic activity and reduced root respiration. However, tolerance and avoidance strategies for drought vary markedly among species. Drought can substantially decrease growth, biomass and photosynthetic activity among seedlings in field and laboratory studies. When compared with the responses to flooding, drought can impose higher seedling mortality and slower growth rates, especially among evergreen species. Results indicate that tolerance and avoidance strategies for drought vary markedly between species. Both seedling recruitment and photosynthetic activity are affected by drought,
For many species, the effects of drought can be as important as flooding for survival and growth, particularly at the seedling phase of establishment, ultimately influencing species composition. In the context of climate change and predicted decreases in precipitation in the Amazon Basin, the effects of drought on plant physiology and species distribution in tropical floodplain forest ecosystems should not be overlooked.
Drought responses; Amazonia; floodplain forests; tree ecology; várzea
A recent lengthening of the growing season in mid and higher latitudes of the northern hemisphere is reported as a clear indicator for climate change impacts. Using data from Germany (1951–2003) and Slovenia (1961–2004), we study whether changes in the start, end, and length of the growing season differ among four deciduous broad-leaved tree species and countries, how the changes are related to temperature changes, and what might be the confounding effects of an insect attack. The functional behaviour of the phenological and climatological time series and their trends are not analysed by linear regression, but by a new Bayesian approach taking into account different models for the functional description (one change-point, linear, constant models). We find advanced leaf unfolding in both countries with the same species order (oak > horse chestnut, beech, and birch). However, this advance is non linear over time and more apparent in Germany with clear change-points in the late 1970s, followed by marked advances (on average 3.67 days decade−1 in the 2000s). In Slovenia, we find a more gradual advance of onset dates (on average 0.8 days decade−1 in the 2000s). Leaf colouring of birch, beech, and oak has been slightly delayed in the last 3 decades, especially in Germany, however with no clear functional behaviour. Abrupt changes in leaf colouring dates of horse chestnut with recent advancing onset dates can be linked across countries to damage by a newly emerging pest, the horse chestnut leaf-miner (Cameraria ohridella). The lengthening of the growing season, more distinct in Germany than in Slovenia (on average 4.2 and 1.0 days decade−1 in the 2000s, respectively), exhibits the same species order in both countries (oak > birch > beech). Damage by horse chestnut leaf-miner leads to reduced lengthening (Germany) and drastic shortening (Slovenia) of the horse chestnut growing season (-12 days decade−1 in the 2000s). Advanced spring leaf unfolding and lengthening of the growing season of oak, beech and birch are highly significantly related to increasing March temperatures in both countries. Only beech and oak leaf unfolding in Germany, which is generally observed later in the year than that of the other two species, is more closely correlated with April temperatures, which comparably exhibit marked change-points at the end of the 1970s.
Growing season; Global warming; Phenology; Deciduous broad-leaved species; Cameraria ohridella; Bayes; Temperature
Hydraulic traits and hydraulic-related structural properties were examined in three deciduous (Hevea brasiliensis, Macaranga denticulate, and Bischofia javanica) and three evergreen (Drypetes indica, Aleurites moluccana, and Codiaeum variegatum) Euphorbiaceae tree species from a seasonally tropical forest in south-western China. Xylem water potential at 50% loss of stem hydraulic conductivity (P50stem) was more negative in the evergreen tree, but leaf water potential at 50% loss of leaf hydraulic conductivity (P50leaf) did not function as P50stem did. Furthermore, P50stem was more negative than P50leaf in the evergreen tree; contrarily, this pattern was not observed in the deciduous tree. Leaf hydraulic conductivity overlapped considerably, but stem hydraulic conductivity diverged between the evergreen and deciduous tree. Correspondingly, structural properties of leaves overlapped substantially; however, structural properties of stem diverged markedly. Consequently, leaf and stem hydraulic traits were closely correlated with leaf and stem structural properties, respectively. Additionally, stem hydraulic efficiency was significantly correlated with stem hydraulic resistance to embolism; nevertheless, such a hydraulic pattern was not found in leaf hydraulics. Thus, these results suggest: (1) that the evergreen and deciduous tree mainly diverge in stem hydraulics, but not in leaf hydraulics, (2) that regardless of leaf or stem, their hydraulic traits result primarily from structural properties, and not from leaf phenology, (3) that leaves are more vulnerable to drought-induced embolism than stem in the evergreen tree, but not always in the deciduous tree and (4) that there exists a trade-off between hydraulic efficiency and safety for stem hydraulics, but not for leaf hydraulics.
Evergreen; Deciduous; Hydraulics; Functional structure
Changes in the timing of phenological events may cause the annual carbon budget of deciduous forests to change. Therefore, one should take such events into account when evaluating the effects of global warming on deciduous forests. In this article, we report on the results of numerical experiments done with a model that includes a phenological module simulating the timing of bud burst and other phenological events and estimating maximum leaf area index.
This study suggests that the negative effects of warming on tree productivity (net primary production) outweigh the positive effects of a prolonged growing season. An increase in air temperature by 3°C (5°C) reduces cumulative net primary production by 21.3% (34.2%). Similarly, cumulative net ecosystem production (the difference between cumulative net primary production and heterotrophic respiration) decreases by 43.5% (64.5%) when temperatures are increased by 3°C (5°C). However, the positive effects of CO2 enrichment (2 × CO2) outweigh the negative effects of warming (<5°C).
Although the model was calibrated and validated for a specific forest ecosystem, the implications of the study may be extrapolated to deciduous forests in cool-temperate zones. These forests share common features, and it can be conjectured that carbon stocks would increase in such forests in the face of doubled CO2 and increased temperatures as long as the increase in temperature does not exceed 5°C.
Background and Aims
Deciduous trees have a seasonal carbon dioxide exchange pattern that is attributed to changes in leaf biochemical properties. However, it is not known if the pattern in leaf biochemical properties – maximum Rubisco carboxylation (Vcmax) and electron transport (Jmax) – differ between species. This study explored whether a general pattern of changes in Vcmax, Jmax, and a standardized soil moisture response accounted for carbon dioxide exchange of deciduous trees throughout the growing season.
The model MAESTRA was used to examine Vcmax and Jmax of leaves of five deciduous trees, Acer rubrum ‘Summer Red’, Betula nigra, Quercus nuttallii, Quercus phellos and Paulownia elongata, and their response to soil moisture. MAESTRA was parameterized using data from in situ measurements on organs. Linking the changes in biochemical properties of leaves to the whole tree, MAESTRA integrated the general pattern in Vcmax and Jmax from gas exchange parameters of leaves with a standardized soil moisture response to describe carbon dioxide exchange throughout the growing season. The model estimates were tested against measurements made on the five species under both irrigated and water-stressed conditions.
Measurements and modelling demonstrate that the seasonal pattern of biochemical activity in leaves and soil moisture response can be parameterized with straightforward general relationships. Over the course of the season, differences in carbon exchange between measured and modelled values were within 6–12 % under well-watered conditions and 2–25 % under water stress conditions. Hence, a generalized seasonal pattern in the leaf-level physiological change of Vcmax and Jmax, and a standardized response to soil moisture was sufficient to parameterize carbon dioxide exchange for large-scale evaluations.
Simplification in parameterization of the seasonal pattern of leaf biochemical activity and soil moisture response of deciduous forest species is demonstrated. This allows reliable modelling of carbon exchange for deciduous trees, thus circumventing the need for extensive gas exchange experiments on different species.
Carbon budget; deciduous trees; modelling; MAESTRA; soil moisture; species response; transpiration; Acer rubrum; Betula nigra; Quercus nuttallii; Q. phellos; Paulownia elongata
Background and Aims
Cambium reactivation after dormancy and budbreak in deciduous trees requires a supply of mobilized reserve materials. The pathway and mode of transfer of these materials are poorly understood.
Transport of reserve materials during cambium reactivation in Populus nigra was investigated by conventional and immunocytochemical TEM analyses, SDS–PAGE, western blotting and intracellular microinjection of fluorescent dyes.
Proteinaceous compounds stored in vacuoles and protein bodies of vascular cells and ray cells disappeared within 3 weeks after cambial reactivation and budbreak. Some of these proteins (32 kDa, 30 kDa and 15 kDa) were labelled by lectin antibodies in SDS–PAGE. The same antibodies were localized to plasmodesmata (PDs) between phloem parenchyma, ray cells and fusiform cambial cells. In addition, proteinaceous particles were localized inside the cytoplasmic sleeves of these PDs during budbreak. During this period, the functional diameter of PDs was about 2·2 nm which corresponds approximately to the Stokes' radius of the detected 15-kDa protein.
Lectin-like reserve proteins or their degradation products seem to be transferred through PDs of phloem parenchyma and rays during cambial reactivation and budbreak. PD transfer of storage proteins is a novelty which supports the concept of symplasmic nutrient supply to the cambial region.
Cambial region; lectins; plasmodesmal trafficking; Populus nigra ‘italica’; size exclusion limit; storage proteins; vascular tissues
Wood formation requires a continuous supply of carbohydrates for structural growth and metabolism. In the montane belt of the central Austrian Alps we monitored the temporal dynamics of xylem growth and non-structural carbohydrates (NSC) in stem sapwood of Pinus sylvestris L. during the growing season 2009, which was characterized by exceptional soil dryness within the study area. Soil water content dropped below 10 % at the time of maximum xylem growth end of May. Histological analyses have been used to describe cambial activity and xylem growth. Determination of NSC was performed using specific enzymatic assays revealing that total NSC ranged from 0.8 to 1.7 % dry matter throughout the year. Significant variations (P < 0.05) of the size of the NSC pool were observed during the growing season. Starch showed persistent abundance throughout the year reaching a maximum shortly before onset of late wood formation in mid-July. Seasonal dynamics of NSC and xylem growth suggest that (i) high sink activity occurred at start of the growing season in spring and during late wood formation in summer and (ii) there was no particular shortage in NSC, which caused P. sylvestris to draw upon stem reserves more heavily during drought in 2009.
1.Insect herbivory can strongly affect ecosystem processes, and its relationship with plant diversity is a central topic in biodiversity–functioning research. However, very little is known about this relationship from complex ecosystems dominated by long-lived individuals, such as forests, especially over gradients of high plant diversity.
2.We analysed insect herbivory on saplings of 10 tree and shrub species across 27 forest stands differing in age and tree species richness in an extraordinarily diverse subtropical forest ecosystem in China. We tested whether plant species richness significantly influences folivory in these highly diverse forests or whether other factors play a more important role at such high levels of phytodiversity.
3.Leaf damage was assessed on 58 297 leaves of 1284 saplings at the end of the rainy season in 2008, together with structural and abiotic stand characteristics.
4.Species-specific mean damage of leaf area ranged from 3% to 16%. Herbivory increased with plant species richness even after accounting for potentially confounding effects of stand characteristics, of which stand age-related aspects most clearly covaried with herbivory. Intraspecific density dependence or other abiotic factors did not significantly influence overall herbivory across forest stands.
5.Synthesis.The positive herbivory–plant diversity relationship indicates that effects related to hypotheses of resource concentration, according to which a reduction in damage by specialized herbivores might be expected as host plant concentration decreases with increasing plant diversity, do not seem to be major determinants for overall herbivory levels in our phytodiverse subtropical forest ecosystem. We discuss the potential role of host specificity of dominant herbivores, which are often expected to show a high degree of specialization in many (sub)tropical forests. In the forest system we studied, a much higher impact of polyphagous species than traditionally assumed might explain the observed patterns, as these species can profit from a broad dietary mix provided by high plant diversity. Further testing is needed to experimentally verify this assumption.
BEF China; biodiversity; ecosystem functioning; Gutianshan; resource concentration; succession; trophic interactions; Zhejiang
The population dynamics of the rubber plantation litter beetle, Luprops tristis Fabricius 1801 (Coleoptera: Tenebrionidae) was assessed in relation to the phenology of leaf shedding and defoliation pattern of para rubber trees, Hevea brasiliensis Müll. Arg (Malpighiales: Euphorbiaceae), during a two year study period. The abundance of adults, larvae and pupae per 1m2 of litter sample was recorded. Post dormancy beetles appeared in leaf litter following annual leaf shedding, whereas larvae, pupae and teneral adults were present after leaf flush. No stages were recorded from plantations following the summer rains until the annual litter fall in the next season. Parental adults peaked at the time of leaf sprouting and tender leaf fall. Larvae and teneral adults peaked at the time of premature fall of green leaves and flowers. Teneral adults of six age classes were recorded and all entered dormancy irrespective of the feeding time available to each age class. Females outnumbered males in the parent generation, while the sex ratio of new generation adults was not biased towards either sex. The phenological stages of rubber trees included leaf fall in late December and early January, leaf sprouting and new leaf production in January and flowering in February. All feeding stages of L. tristis peaked in abundance when premature leaves are most abundant in the leaf litter. Prediction of the timing of appearance of various developmental stages of L. tristis in plantations, invasion into buildings and intensity of population build up in rubber belts is possible by tracking the phenology of leaf fall in rubber plantations, time of return of post dormancy adults and the onset of summer rainfall. Perfect synchrony was recorded between the field return of parental adults with annual leaf shedding, the oviposition phase of parental adults with tender leaf fall at the time of leaf sprouting, and larval and teneral adult stages with premature fall of leaves. Premature leaf availability is suggested as contributing to the reproductive efficiency of parental adults, the survival of early developmental stages and of new generation adults during dormancy.
Mupli beetles; aggregation; Hevea brasiliensis RRII 105
Background and Aims
Oil palm flowering and fruit production show seasonal maxima whose causes are unknown. Drought periods confound these rhythms, making it difficult to analyse or predict dynamics of production. The present work aims to analyse phenological and growth responses of adult oil palms to seasonal and inter-annual climatic variability.
Two oil palm genotypes planted in a replicated design at two sites in Indonesia underwent monthly observations during 22 months in 2006–2008. Measurements included growth of vegetative and reproductive organs, morphology and phenology. Drought was estimated from climatic water balance (rainfall – potential evapotranspiration) and simulated fraction of transpirable soil water. Production history of the same plants for 2001–2005 was used for inter-annual analyses.
Drought was absent at the equatorial Kandista site (0°55′N) but the Batu Mulia site (3°12′S) had a dry season with variable severity. Vegetative growth and leaf appearance rate fluctuated with drought level. Yield of fruit, a function of the number of female inflorescences produced, was negatively correlated with photoperiod at Kandista. Dual annual maxima were observed supporting a recent theory of circadian control. The photoperiod-sensitive phases were estimated at 9 (or 9 + 12 × n) months before bunch maturity for a given phytomer. The main sensitive phase for drought effects was estimated at 29 months before bunch maturity, presumably associated with inflorescence sex determination.
It is assumed that seasonal peaks of flowering in oil palm are controlled even near the equator by photoperiod response within a phytomer. These patterns are confounded with drought effects that affect flowering (yield) with long time-lag. Resulting dynamics are complex, but if the present results are confirmed it will be possible to predict them with models.
Photoperiodism; Elaeis guineensis; flowering; phyllochron; drought; radiation use efficiency; sink–source relationships; phenotypic plasticity
Within a dry inner Alpine valley in the Eastern Central Alps (750 m a.s.l., Tyrol, Austria) the influence of climate variables (precipitation, air humidity, temperature) and soil water content on intra-annual dynamics of tree-ring development was determined in Scots pine (Pinus sylvestris L.) at two sites differing in soil water availability (xeric and dry-mesic site). Radial stem development was continuously followed during 2007 and 2008 by band dendrometers and repeated micro-sampling of the developing tree rings of mature trees. Daily and seasonal fluctuations of the stem radius, which reached almost half of total annual increment, primarily reflected changes in tree water status and masked radial stem growth especially during drought periods in spring. However, temporal dynamics of intra-annual radial growth determined by both methods were found to be quite similar, when onset of radial growth in dendrometer traces was defined by the occurrence of first enlarging xylem cells. Radial increments during the growing period, which lasted from early April through early August showed statistically significant relationships with precipitation (Kendall τ = 0.234, p < 0.01, and τ = 0.184, p < 0.05, at the xeric and dry-mesic site, respectively) and relative air humidity (Pearson r = 0.290, p < 0.05, and r = 0.306, p < 0.05 at the xeric and dry-mesic site, respectively). Soil water content and air temperature had no influence on radial stem increment. Culmination of radial stem growth was detected at both study plots around mid-May, prior to occurrence of more favourable climatic conditions, i.e. an increase in precipitation during summer. We suggest that the early decrease in radial growth rate is due to a high belowground demand for carbohydrates to ensure adequate resource acquisition on the drought prone substrate.
Dendrometer; Drought; Dry inner Alpine valley; Pinus sylvestris; Radial growth; Xylem cell analysis
In Amazonian floodplain forests, >1000 tree species grow in an environment subject to extended annual submergence which can last up to 9 months each year. Water depth can reach 10 m, fully submerging young and also adult trees, most of which reproduce during the flood season. Complete submergence occurs regularly at the seedling or sapling stage for many species that colonize low-lying positions in the flooding gradient. Here hypoxic conditions prevail close to the water surface in moving water, while anaerobic conditions are common in stagnant pools. Light intensities in the floodwater are very low.
Questions and Aims
Despite a lack of both oxygen and light imposed by submergence for several months, most leafed seedlings survive. Furthermore, underwater growth has also been observed in several species in the field and under experimental conditions. The present article assesses how these remarkable plants react to submergence and discusses physiological mechanisms and anatomical adaptations that may explain their success.
Adaptation; Amazonian floodplains; darkness; environmental stress; flooding; hypoxia; submergence tolerance; trees; underwater photosynthesis; woody species
Olive trees are a classic component of Mediterranean environments and some of them are known historically to be very old. In order to evaluate the possibility to use olive tree-rings for dendrochronology, we examined by various methods the reliability of olive tree-rings identification. Dendrochronological analyses of olive trees growing on the Aegean island Santorini (Greece) show that the determination of the number of tree-rings is impossible because of intra-annual wood density fluctuations, variability in tree-ring boundary structure, and restriction of its cambial activity to shifting sectors of the circumference, causing the tree-ring sequences along radii of the same cross section to differ.
Lianas are an important component of most tropical forests, where they vary in abundance from high in seasonal forests to low in aseasonal forests. We tested the hypothesis that the physiological ability of lianas to fix carbon (and thus grow) during seasonal drought may confer a distinct advantage in seasonal tropical forests, which may explain pan-tropical liana distributions. We compared a range of leaf-level physiological attributes of 18 co-occurring liana and 16 tree species during the wet and dry seasons in a tropical seasonal forest in Xishuangbanna, China. We found that, during the wet season, lianas had significantly higher CO2 assimilation per unit mass (Amass), nitrogen concentration (Nmass), and δ13C values, and lower leaf mass per unit area (LMA) than trees, indicating that lianas have higher assimilation rates per unit leaf mass and higher integrated water-use efficiency (WUE), but lower leaf structural investments. Seasonal variation in CO2 assimilation per unit area (Aarea), phosphorus concentration per unit mass (Pmass), and photosynthetic N-use efficiency (PNUE), however, was significantly lower in lianas than in trees. For instance, mean tree Aarea decreased by 30.1% from wet to dry season, compared with only 12.8% for lianas. In contrast, from the wet to dry season mean liana δ13C increased four times more than tree δ13C, with no reduction in PNUE, whereas trees had a significant reduction in PNUE. Lianas had higher Amass than trees throughout the year, regardless of season. Collectively, our findings indicate that lianas fix more carbon and use water and nitrogen more efficiently than trees, particularly during seasonal drought, which may confer a competitive advantage to lianas during the dry season, and thus may explain their high relative abundance in seasonal tropical forests.
Liana distribution; Nitrogen-use efficiency; Tropical forest physiology; Water-use efficiency
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.
Plant functional types; Leaf plasticity; Shade acclimation; Evergreen leaves; Leaf mass-per-area