Response Of Woody Plants Research Articles

Genome-wide transcriptional and physiological responses to drought stress in leaves and roots of two willow genotypes.

BackgroundDrought is a major environmental stress that can have severe impacts on plant productivity and survival. Understanding molecular mechanisms of drought responses is crucial in order to breed for drought adapted plant cultivars. The aim of the present study was to investigate phenotypic and transcriptional drought responses in two willow genotypes (520 and 592) originating from an experimental cross between S. viminalis × (S. viminalis × S. schwerinii). Willows are woody perennials in the Salicaceae plant family that are grown as bioenergy crops worldwide.MethodsAn experiment was conducted where plants were exposed to drought and different eco-physiological parameters were assessed. RNA-seq data was furthermore generated with the Illumina technology from root tips and leaves from plants grown in drought and well-watered (WW) conditions. The RNA-seq data was assembled de novo with the Trinity assembler to create a reference gene set to which the reads were mapped in order to obtain differentially expressed genes (DEGs) between the drought and WW conditions. To investigate molecular mechanisms involved in the drought response, GO enrichment analyses were conducted. Candidate genes with a putative function in the drought response were also identified.ResultsA total of 52,599 gene models were obtained and after filtering on gene expression (FPKM ≥ 1), 35,733 gene models remained, of which 24,421 contained open reading frames. A total of 5,112 unique DEGs were identified between drought and WW conditions, of which the majority were found in the root tips. Phenotypically, genotype 592 displayed less growth reduction in response to drought compared to genotype 520. At the transcriptional level, genotype 520 displayed a greater response in the leaves as more DEGs were found in genotype 520 compared to genotype 592. In contrast, the transcriptional responses in the root tips were rather similar between the two genotypes. A core set of candidate genes encoding proteins with a putative function in drought response was identified, for example MYBs and bZIPs as well as chlorophyll a/b binding proteins.DiscussionWe found substantial differences in drought responses between the genotypes, both at the phenotypic and transcriptional levels. In addition to the genotypic variation in several traits, we also found indications for genotypic variation in trait plasticity, which could play a role in drought adaptation. Furthermore, the two genotypes displayed overall similar transcriptional responses in the root tips, but more variation in the leaves. It is thus possible that the observed phenotypic differences could be a result of transcriptional differences mostly at the leaf level.ConclusionsThis study has contributed to a better general understanding of drought responses in woody plants, specifically in willows, and has implications for breeding research towards more drought adapted plants.Electronic supplementary materialThe online version of this article (doi:10.1186/s12870-015-0630-2) contains supplementary material, which is available to authorized users.

Does the growth response of woody plants to elevated CO2 increase with temperature? A model-oriented meta-analysis.

The temperature dependence of the reaction kinetics of the Rubisco enzyme implies that, at the level of a chloroplast, the response of photosynthesis to rising atmospheric CO2 concentration (Ca ) will increase with increasing air temperature. Vegetation models incorporating this interaction predict that the response of net primary productivity (NPP) to elevated CO2 (eCa ) will increase with rising temperature and will be substantially larger in warm tropical forests than in cold boreal forests. We tested these model predictions against evidence from eCa experiments by carrying out two meta-analyses. Firstly, we tested for an interaction effect on growth responses in factorial eCa ×temperature experiments. This analysis showed a positive, but nonsignificant interaction effect (95% CI for above-ground biomass response=-0.8, 18.0%) between eCa and temperature. Secondly, we tested field-based eCa experiments on woody plants across the globe for a relationship between the eCa effect on plant biomass and mean annual temperature (MAT). This second analysis showed a positive but nonsignificant correlation between the eCa response and MAT. The magnitude of the interactions between CO2 and temperature found in both meta-analyses were consistent with model predictions, even though both analyses gave nonsignificant results. Thus, we conclude that it is not possible to distinguish between the competing hypotheses of no interaction vs. an interaction based on Rubisco kinetics from the available experimental database. Experiments in a wider range of temperature zones are required. Until such experimental data are available, model predictions should aim to incorporate uncertainty about this interaction.

Xylem parenchyma cell walls lack a gravitropic response in conifer compression wood.

Cell wall fluorescence and immunocytochemistry demonstrate that xylem parenchyma cell walls do not show changes in structure and composition related to gravitropic response comparable to those of tracheids, even when they have lignified secondary cell walls. Tracheid cell walls in compression wood have altered composition and structure which generates the strain responsible for correction of stem lean as part of the gravitropic response of woody plants. Xylem parenchyma cell walls vary among conifer species and can be lignified secondary walls (spruce) or unlignified primary walls (pine). It can be expected that xylem parenchyma with lignified secondary cell walls might show features of compression wood comparable to those of tracheids that have a similar type of cell wall. A comparison of xylem parenchyma cell walls in normal and compression wood in species with lignified and non-lignified parenchyma cell walls provides a unique opportunity to understand the process of reaction wood formation in conifers. Using both UV/visible fluorescence microscopy of cell wall fluorophores and immunocytochemistry of galactan and mannan epitopes, we demonstrate that xylem parenchyma cell walls do not show the changes in composition and structure typical of compression wood tracheids. Adjacent cells of different types but with similar cell wall structure can undergo cell wall developmental changes related to support or defence functions independent of their neighbours. Tracheids are sensitive to gravitropic signals while xylem parenchyma cells are not.

Biochar and forest restoration: a review and meta-analysis of tree growth responses

“Biochar”, or charcoal intended for use as a soil amendment, has received great attention in recent years as a means of enhancing carbon sequestration and soil properties in agricultural systems. Here we address the potential for biochar use in the context of forest restoration, reviewing relevant experimental studies on biochar use in forest ecosystems, the properties of chars generated from wood waste material, and available data on tree growth responses to biochar. To our knowledge the earliest mention of char use as a soil amendment is actually specifically in the context of forest restoration (in the 1820s in Scotland). Wood waste biochars have an unusual set of properties that suggest their applicability in a forest restoration context: namely, high recalcitrance promoting long-lasting effects, retention of cations, anions, and water, in the soil, sorptive properties that can reduce bioavailability of a wide range of toxic materials, and relative ease of production from locally available feedstocks. A meta-analysis of recent studies on biochar responses of woody plants indicates a potential for large tree growth responses to biochar additions, with a mean 41 % increase in biomass. Responses are especially pronounced at early growth stages, and appear to be higher in boreal and tropical than in temperate systems, and in angiosperms than conifers; however, there is high variability, and field studies are few. The properties of biochars also vary greatly depending on feedstock and pyrolysis conditions; while this complicates their use, it provides a means to design biochars for specific restoration situations and objectives. We conclude that there is great promise for biochar to play an important role in a wide variety of forest restoration efforts, specifically as a replacement product for other forms of organic matter and liming agents.

INTEGRATED ASSESSMENT AND GISMAPPING OF THE ENVIRONMENTAL STATE OF THE CITY OF VORONEZH (RUSSIA)

The authors have created a geoinformation-analytical system (GIS) for integrated assessment and mapping of the ecological conditions of the territory according to the criteria of anthropogenic impact and quality of the urban environment, as well as the response of woody plants and the health of the child population (on the example of Voronezh – the largest industrial city of the Central Chernozem region). It has been identified that anthropogenic pollution is formed by the industrial-transport sector and varies with regard to the features of the functional planning infrastructure; near the industrial facilities of the petrochemical profile in the left-Bank sector of the city, conditions for the existence of woody plants significantly worsen, which is manifested in the inhibition of their development; child morbidity rate is significantly higher in industrially polluted neighborhoods with high load of pollutant emissions from industry and transport. The diseases primarily associated with pollution are congenital anomalies, neoplasms, endocrine pathology and diseases of the urogenital area. The industrial zone is the main contributor to the total pollution of air, but the transport zone is the main contributor to the total pollution of soil and snow cover.

Proteomic analysis reveals differences in tolerance to acid rain in two broad-leaf tree species, Liquidambar formosana and Schima superba.

Acid rain (AR) is a serious environmental issue inducing harmful impacts on plant growth and development. It has been reported that Liquidambar formosana, considered as an AR-sensitive tree species, was largely injured by AR, compared with Schima superba, an AR-tolerant tree species. To clarify the different responses of these two species to AR, a comparative proteomic analysis was conducted in this study. More than 1000 protein spots were reproducibly detected on two-dimensional electrophoresis gels. Among them, 74 protein spots from L. formosana gels and 34 protein spots from S. superba gels showed significant changes in their abundances under AR stress. In both L. formosana and S. superba, the majority proteins with more than 2 fold changes were involved in photosynthesis and energy production, followed by material metabolism, stress and defense, transcription, post-translational and modification, and signal transduction. In contrast with L. formosana, no hormone response-related protein was found in S. superba. Moreover, the changes of proteins involved in photosynthesis, starch synthesis, and translation were distinctly different between L. formosana and S. superba. Protein expression analysis of three proteins (ribulose-1,5-bisphosphate carboxylase/oxygenase large subunit, ascorbate peroxidase and glutathione-S-transferase) by Western blot was well correlated with the results of proteomics. In conclusion, our study provides new insights into AR stress responses in woody plants and clarifies the differences in strategies to cope with AR between L. formosana and S. superba.

Comparing bark thickness: testing methods with bark–stem data from two South African fire‐prone biomes

AbstractAimsBark thickness–stem diameter relationships are non‐linear above a stem diameter threshold in many woody species, which makes relative bark thickness measures dependent on the range of stem diameters that are sampled. This influences the appropriateness of different methods for comparing fire responses of woody plants across studies. Here we develop a framework for bark thickness comparisons by evaluating relative bark thickness estimates and bark thickness predictions obtained from linear and curved models fitted to raw and log‐transformed bark–stem data. We use this framework to contrast bark thickness among fynbos and savanna plant functional groups.LocationFynbos (17 species) and savanna (21 species) systems in South Africa.MethodsThe linear subset of bark–stem data was identified using a three‐step procedure. Linear regressions (with and without an intercept) and curved models (allometric and modified exponential models) were fitted to the linear subset and complete raw bark–stem data set, respectively. In addition, linear regression models were fitted to the log‐transformed complete bark–stem data set. Regression slopes and bark thickness predictions obtained from these different approaches were compared, to determine which method provides the most robust metric for comparing bark thickness. Bark thickness was compared among fynbos resprouter guilds and acacias from low‐ and high‐fire savannas.ResultsThe slope of the regression model fitted to the linear subset of raw bark–stem data provides a reliable metric for general comparisons of relative bark thickness. Bark thickness predictions from the curved and log models were comparable at 20‐cm stem diameter, but the log model underestimated bark thickness at 5 cm for certain species. Relative bark thickness was significantly higher in: (1) fynbos fire resisters and epicormic resprouters than in non‐resprouters; and (2) acacias from high‐ vs low‐fire savannas.ConclusionsThe slope of the regression model fitted to the linear subset of raw bark–stem data is a useful metric for bark thickness comparisons across studies, and compares favourably with bark thickness predictions derived from models fitted to the complete bark–stem data set. Fynbos and savanna trends support the proposition that relative bark thickness reflects differences in woody plant responses to fire and indicate the modal fire regime of ecosystems.

Effects of Growth Form and Functional Traits on Response of Woody Plants to Clearing and Fragmentation of Subtropical Rainforest

The conservation implications of large-scale rainforest clearing and fragmentation on the persistence of functional and taxonomic diversity remain poorly understood. If traits represent adaptive strategies of plant species to particular circumstances, the expectation is that the effect of forest clearing and fragmentation will be affected by species functional traits, particularly those related to dispersal. We used species occurrence data for woody plants in 46 rainforest patches across 75,000 ha largely cleared of forest by the early 1900s to determine the combined effects of area reduction, fragmentation, and patch size on the taxonomic structure and functional diversity of subtropical rainforest. We compiled species trait values for leaf area, seed dry mass, wood density, and maximum height and calculated species niche breadths. Taxonomic structure, trait values (means, ranges), and the functional diversity of assemblages of climbing and free-standing plants in remnant patches were quantified. Larger rainforest patches had higher species richness. Species in smaller patches were taxonomically less related than species in larger patches. Free-standing plants had a high percentage of frugivore dispersed seeds; climbers had a high proportion of small wind-dispersed seeds. Connections between the patchy spatial distribution of free-standing species, larger seed sizes, and dispersal syndrome were weak. Assemblages of free-standing plants in patches showed more taxonomic and spatial structuring than climbing plants. Smaller isolated patches retained relatively high functional diversity and similar taxonomic structure to larger tracts of forest despite lower species richness. The response of woody plants to clearing and fragmentation of subtropical rainforest differed between climbers and slow-growing mature-phase forest trees but not between climbers and pioneer trees. Quantifying taxonomic structure and functional diversity provides an improved basis for conservation planning and management by elucidating the effects of forest-area reduction and fragmentation. Efectos de la Forma de Crecimiento y Atributos Funcionales en la Respuesta de Plantas Leñosas al Desmonte y Fragmentación de Bosque Lluvioso Subtropical.

Transpirational demand affects aquaporin expression in poplar roots

Isohydric plants tend to maintain a water potential homeostasis primarily by controlling water loss via stomatal conductance. However, there is accumulating evidence that plants can also modulate water uptake in a dynamic manner. The dynamics of water uptake are influenced by aquaporin-mediated changes in root hydraulics. Most studies in this area have been conducted on herbaceous plants, and less is known about responses of woody plants. Here a study was conducted to determine how roots of hybrid poplar plants (Populus trichocarpa×deltoides) respond to a step change in transpirational demand. The main objective was to measure the expression of selected aquaporin genes and to assess how transcriptional responses correspond to changes in root water flow (Q R) and other parameters of water relations. A subset of plants was grown in shade and was subsequently exposed to a 5-fold increase in light level. Another group of plants was grown at ~95% relative humidity (RH) and was then subjected to lower RH while the light level remained unchanged. Both plant groups experienced a transient drop in stem water potentials. At 28h after the increase in transpirational demand, water potentials recovered. This recovery was associated with changes in the expression of PIP1 and PIP2 subfamily genes and an increase in Q R. Stomata of plants growing at high RH were larger and showed incomplete closure after application of abscisic acid. Since stomatal conductance remained high and unchanged in these plants, it is suggested that the recovery in water potential in these plants was largely driven by the increase in Q R.

Global identification of miRNAs and targets in Populus euphratica under salt stress

Populus euphratica, a typical hydro-halophyte, is ideal for studying salt stress responses in woody plants. MicroRNAs (miRNAs) are endogenous non-coding small RNAs that fulfilled an important post-transcriptional regulatory function. MiRNA may regulate tolerance to salt stress but this has not been widely studied in P. euphratica. In this investigation, the small RNAome, degradome and transcriptome were studied in salt stress treated P. euphratica by deep sequencing. Two hundred and eleven conserved miRNAs between Populus trichocarpa and P. euphratica have been found. In addition, 162 new miRNAs, belonging to 93 families, were identified in P. euphratica. Degradome sequencing experimentally verified 112 targets that belonged to 51 identified miRNAs, few of which were known previously in P. euphratica. Transcriptome profiling showed that expression of 15 miRNA-target pairs displayed reverse changing pattern under salt stress. Together, these results indicate that, in P. euphratica under salt stress, a large number of new miRNAs could be discovered, and both known and new miRNA were functionally cleaving to their target mRNA. Expression of miRNA and target were correspondingly induced by salt stress but that it was a complex process in P. euphratica.

Effects of salt stress on ion content, antioxidant enzymes and protein profile in different tissues of Broussonetia papyrifera

Although some plant responses to salinity have been characterized, the precise mechanisms by which salt stress damages plants are still poorly understood especially in woody plants. In the present study, the physiological and biochemical responses of Broussonetia papyrifera, a tree species of the family, Moraceae, to salinity were studied. In vitro-produced plantlets of B. papyrifera were treated with varying levels of NaCl (0, 50, 100 and 150mM) in hydroponic culture. Changes in ion contents, accumulation of H2O2, as well as the activities and isoform profiles of superoxide dismutase (SOD), peroxidase (POD) and catalase (CAT) in the leaves, stems and roots were investigated. Under salt stress, there was higher Na+ accumulation in roots than in stems and leaves, and Ca2+, Mg2+ and P3+ content, as well as K+/Na+ ratio were affected. NaCl treatment induced an increase in H2O2 contents in the tissues of B. papyrifera. The work demonstrated that activities of antioxidant defense enzymes changed in parallel with the increased H2O2 and salinity appeared to be associated with differential regulation of distinct SOD and POD isoenzymes. Moreover, SDS-PAGE analysis of total proteins extracted from leaves and roots of control and NaCl-treated plantlets revealed that in the leaves salt stress was associated with decrease or disappearance of some protein bands, and induction of a new protein band after exposure to 100 and 150mM NaCl. In contrast, NaCl stress had little effect on the protein pattern in the roots. In summary, these findings may provide insight into the mechanisms of the response of woody plants to salt stress.

Fire persistence traits can be used to predict vegetation response to changing fire regimes at expansive landscape scales – an Australian example

AbstractAim Building on a substantial literature addressing the fire responses of woody plants, particularly under mediterranean climates, we assess the extent to which fire persistence traits can be used to predict vegetation responses to fire regime changes in fire‐prone arid and savanna landscape settings.Location Australia, applying data from arid central to monsoonal northern regions (11–26° S, 129–138° E).Methods With reference to a substantial sub‐continental floristics dataset, we first assigned the fire response (obligate seeder, resprouter) and seedbank persistence (transient, dormant) of rapid and longer‐maturing (> 3 years) woody taxa. Using logistic regression, we then modelled the proportions of taxa possessing these traits as a function of mean annual rainfall (highly correlated with fire frequency) and terrain roughness (a measure of topographic variability) in 0.25° × 0.25° and 1° × 1° grid cells. Separate assessments were undertaken with datasets for 1264 sclerophyll and 236 rain forest taxa.Results This woody flora is characterized by taxa exhibiting mostly resprouting and dormant seedbank traits that promote site persistence. While numbers of obligate seeder and resprouter taxa were related positively to both rainfall and roughness, the relative abundance of both sclerophyll and rain forest obligate seeders decreased significantly with rainfall. The relative abundance of sclerophyll (especially long‐lived) obligate seeders alone increased with topographic roughness. The proportion of taxa with transient seedbanks increased with rainfall in resprouters generally, and in rain forest obligate seeders alone.Main conclusions We find that resprouters are favoured on more productive, fire‐prone sites, and obligate seeders are favoured in less productive, more fire‐protected settings. Seedbank persistence responses are more variable. These findings concur generally with theoretical constructs, and support comparable assessments in Australian and other fire‐prone systems ranging from mediterranean to boreal environments. Our observations illustrate that resprouting and obligate seeding syndromes, but not necessarily seedbank persistence, are useful predictors of vegetation responses to changing fire regime conditions at large landscape scales.

Irrigation, organic fertilization and species successional stage modulate the response of woody seedlings to herbaceous competition in a semi‐arid quarry restoration

AbstractQuestionIs the response of woody plants to herbaceous competition affected by the level of irrigation and organic fertilization? Does the magnitude and the sign of this response depend on the successional status of woody species?LocationA limestone quarry in a semi‐arid Mediterranean area inSE Spain.MethodsWe studied seedling survival and growth of six woody species with contrasting ecological strategies (early‐successional vs late‐successional) and their associated herbaceous layer under different irrigation and organic fertilization levelsResultsElevated water supply, but not organic fertilization, increased herbaceous cover. The response of woody seedlings and changes in herbaceous cover to a contrasting resource supply was species‐specific. Seedling survival and growth of late‐successional species increased at high levels of irrigation. Pioneer species showed the opposite trend, suggesting that increases in water availability may outweigh the deleterious effect of increased herbaceous competition in late‐successional species but not in early‐successional species.ConclusionsOur study provides further insights into the role of techniques such as irrigation and organic fertilization, which are commonly used in conjunction with hydroseeding, in defining woody plant establishment and successional trajectories in semi‐arid areas under restoration. Our results will contribute to improve restoration success by reconciling two major objectives of restoration: soil protection and secondary succession. We recommend a new approach for the restoration of limestone quarries. This approach comprises the planting of late‐successional species on randomly distributed nutrient‐rich patches of high availability of water and nutrients, and the use of pioneer species in the surrounding nutrient poor and drier matrix. This approach should promote optimal ecosystem functioning, including soil protection, while increasing biodiversity and fostering secondary succession.

Responses to environmental stresses in woody plants: key to survive and longevity

Environmental stresses have adverse effects on plant growth and productivity, and are predicted to become more severe and widespread in decades to come. Especially, prolonged and repeated severe stresses affecting growth and development would bring down long-lasting effects in woody plants as a result of its long-term growth period. To counteract these effects, trees have evolved specific mechanisms for acclimation and tolerance to environmental stresses. Plant growth and development are regulated by the integration of many environmental and endogenous signals including plant hormones. Acclimation of land plants to environmental stresses is controlled by molecular cascades, also involving cross-talk with other stresses and plant hormone signaling mechanisms. This review focuses on recent studies on molecular mechanisms of abiotic stress responses in woody plants, functions of plant hormones in wood formation, and the interconnection of cell wall biosynthesis and the mechanisms shown above. Understanding of these mechanisms in depth should shed light on the factors for improvement of woody plants to overcome severe environmental stress conditions.

Quantitative comparison of chemical, biological and mechanical induction of secondary compounds in Pinus pinaster seedlings

Chemical elicitors and mechanical treatments simulating real insect herbivory have been increasingly used to study induced defensive responses in woody plants. However, simultaneous quantitative comparisons of plant chemical defences elicited by real and simulated herbivory have received little attention. In this paper we compared the effects of real herbivory, simulated herbivory using two chemical elicitors, and mechanical damage treatments on the quantitative secondary chemistry of Pinus pinaster juveniles (namely on non-volatile resin in the stem and total phenolics in the needles). The real herbivory involved phloem wounding by Hylobius abietis and defoliation by Brachyderes lusitanicus (two pine weevils); the chemical elicitors to simulate herbivory induction were 40 mM methyl jasmonate (MJ) and 20 μM benzothiadiazole (BTH); and the mechanical treatments included phloem wounding and needle clipping. We also performed an additional experiment for assessing at what extent insect extracts could increase plant responses over mechanical damage. Chemical induction with MJ, mechanical wounding and real phloem herbivory by H. abietis all produced quantitatively similar results, increasing the concentration of resin in the stem and total phenolics in the needles by equivalent magnitudes. Exogenous application of BTH increased the concentration of phenolic compounds in pine needles, but did not show the same effect on stem resin. Contrastingly, we did not find significant changes in the concentration of resin in the stem or phenolics in the needles after needle clipping and B. lusitanicus feeding. Mechanical damage followed by the application of extracts from the insects B. lusitanicus and H. abietis on the injured tissues did not increase the responses in comparison to mechanical damage alone. The fact that strong induced responses elicited by phloem wounding insects are equally elicited by phloem injuries suggests that defences in pine trees are raised with low specificity regarding biotic enemies. Results from this paper support future methodological approaches using chemical elicitors and mechanical damage as simulated herbivory treatments for the experimental induction of conifer defences.

Woody plants and the prediction of climate-change impacts on bird diversity

Current methods of assessing climate-induced shifts of species distributions rarely account for species interactions and usually ignore potential differences in response times of interacting taxa to climate change. Here, we used species-richness data from 1005 breeding bird and 1417 woody plant species in Kenya and employed model-averaged coefficients from regression models and median climatic forecasts assembled across 15 climate-change scenarios to predict bird species richness under climate change. Forecasts assuming an instantaneous response of woody plants and birds to climate change suggested increases in future bird species richness across most of Kenya whereas forecasts assuming strongly lagged woody plant responses to climate change indicated a reversed trend, i.e. reduced bird species richness. Uncertainties in predictions of future bird species richness were geographically structured, mainly owing to uncertainties in projected precipitation changes. We conclude that assessments of future species responses to climate change are very sensitive to current uncertainties in regional climate-change projections, and to the inclusion or not of time-lagged interacting taxa. We expect even stronger effects for more specialized plant-animal associations. Given the slow response time of woody plant distributions to climate change, current estimates of future biodiversity of many animal taxa may be both biased and too optimistic.

Moderate soil compaction: Implications on growth and architecture in seedlings of 17 woody plant species

Generally, soil compaction is a stress factor affecting negatively the plant growth, but its effects vary between species and with the soil compaction range. The objective of this study is to know the different growth responses of 17 woody species subjected to moderate soil compaction, because most of the studies about this subject compare the effects in treatments with a wide and discrete compaction range. We explore the effects of moderate soil compaction on a continuous scale (0.1–1.0 MPa) on seedling growth. Seedlings of 17 woody species (deciduous and evergreens) mainly from Mediterranean ecosystems were grown in near optimal conditions (light, temperature and water) in a greenhouse with a sandy substrate. In general, there was a great variability of the responses depending on the species and the studied variable. About 53% of the species showed a higher total biomass with a moderate increase in soil compaction possibly being due to a greater root–soil contact. In the same way, 41% of species increase the relative growth rate and 35% the total area. Nevertheless, in spite of these positive effects on growth, for some species (23%) there was a decrease in the root proportion with soil resistance, as result of soil strength. These effects of moderate soil compaction could sum up in two general responses of woody plants: growth increment and architecture distortion. This might be relevant as a lower root investment may be a disadvantage under drought conditions. Finally, a simple conceptual model is proposed to understand the general effects of soil compaction on growth and biomass allocation.

The effect of prolonged flooding on the bark of mangrove trees

Growth and physiological response of woody plants to flooding have been analyzed in detail; however, relatively few studies have been oriented towards the effects of water immersion on cambial activity and wood and bark anatomy of trees that are growing in prolonged flooding conditions. These studies are important to understand the possible effects of predicted sea level rising in mangroves as a consequence of global warming. We studied five species growing in a mangrove forest, sampling three to six trees of each species, in sites that have the longest flooding period. Differences in bark appearance and phloem structure between the submerged stem portion and the portion of the stem above the water surface exist in all species. Although aerenchyma formation and stem hypertrophy are the most common events related to flooding, each type of tissue responded differently. Annona glabra L., Laguncularia racemosa (L.) Gaertn f. and Hibiscus tiliaceus L. developed rythidome. Avicennia germinans (L.) Stearn developed rythidome only in the submerged stem portion. Phyllanthus elsiae Urb., developed one periderm in both stem portions. Species that developed rythidome also developed aerenchyma between periderms and in the phellem. H. tiliaceus and P. elsiae, showed the highest values for anatomical phloem and periderm characters below water surface, while an inverse tendency was observed in A. glabra and L. racemosa, suggesting that prolonged flooding modifies vascular cambium and phellogen differently. Results indicate that sea level rising would affect distribution of the species according to their specific flooding tolerance.

Effect of Mn toxicity on morphological and physiological changes in two Populus cathayana populations originating from different habitats

The cuttings of Populus cathayana were exposed to four different manganese (Mn) concentrations (0, 0.1, 0.5 and 1 mM) in a greenhouse to investigate the toxicity of Mn and the detoxifying responses of woody plants. Two contrasting populations of P. cathayana, which were from wet and dry climate regions in western China, respectively, were examined in our study. The results showed that high concentration of Mn caused significant decrease in shoot height, biomass accumulation, and leaf number and leaf areas. Injuries to the anatomical features of leaves were also found as the reduced thickness of palisade and spongy parenchyma, the decreased density in the conducting tissue and the collapse and split in the meristematic tissue in the central vein. Moreover, Mn treatments caused the accumulation of hydrogen peroxide (H2O2), and then resulted in oxidative stress indicated by the oxidation of proteins and DNA. Many physiological responses were employed to cope with the toxicity of Mn, including the increase in the contents of non-protein thiol (NP-SH), phytochelatins (PCs) and phenolics compounds and the stimulated activities of guaiacol peroxidase (GPX) and polyphenol oxidase (PPO) for the chelation of Mn and for the antioxidation of reactive oxygen species. The population from dry climate habitat showed a lower leaf concentration of Mn, higher contents of the chelators, and higher activities of GPX and PPO than did the wet climate population at the same Mn treatment, thereby possessing a superior Mn tolerance. In both populations, most of the Mn was accumulated in the shoot, which is favorable regarding phytoremediation.

EFFECTS OF FOUR DECADES OF FIRE MANIPULATION ON WOODY VEGETATION STRUCTURE IN SAVANNA

The amount of carbon stored in savannas represents a significant uncertainty in global carbon budgets, primarily because fire causes actual biomass to differ from potential biomass. We analyzed the structural response of woody plants to long-term experimental burning in savannas. The experiment uses a randomized block design to examine fire exclusion and the season and frequency of burn in 192 7-ha experimental plots located in four different savanna ecosystems. Although previous studies would lead us to expect tree density to respond to the fire regime, our results, obtained from four different savanna ecosystems, suggest that the density of woody individuals was unresponsive to fire. The relative dominance of small trees was, however, highly responsive to fire regime. The observed shift in the structure of tree populations has potentially large impacts on the carbon balance. However, the response of tree biomass to fire of the different savannas studied were different, making it difficult to generalize about the extent to which fire can be used to manipulate carbon sequestration in savannas. This study provides evidence that savannas are demographically resilient to fire, but structurally responsive.