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PubMed2003

Response of C3 and C4 plants to middle-Holocene climatic variation near the prairie-forest ecotone of Minnesota.

Paleorecords of the middle Holocene (MH) from the North American mid-continent can offer insights into ecological responses to pervasive drought that may accompany future climatic warming. We analyzed MH sediments from West Olaf Lake (WOL) and Steel Lake (SL) in Minnesota to examine the effects of warm/dry climatic conditions on prairie-woodland ecosystems. Mineral composition and carbonate delta(18)O were used to determine climatic variations, whereas pollen assemblages, charcoal delta(13)C, and charcoal accumulation rates were used to reconstruct vegetation composition, C(3) and C(4) plant abundance, and fire. The ratio of aragonite/calcite at WOL and delta(18)O at SL suggest that pronounced droughts occurred during the MH but that drought severity decreased with time. From charcoal delta(13)C data we estimated that the MH abundance of C(4) plants averaged 50% at WOL and 43% at SL. At WOL C(4) abundance was negatively correlated with aragonite/calcite, suggesting that severe moisture deficits suppressed C(4) plants in favor of weedy C(3) plants (e.g., Ambrosia). As climate ameliorated C(4) abundance increased (from approximately 33 to 66%) at the expense of weedy species, enhancing fuel availability and fire occurrence. In contrast, farther east at SL where climate was cooler and wetter, C(4) abundance showed no correlation with delta(18)O-inferred aridity. Woody C(3) plants (e.g., Quercus) were more abundant, biomass flammability was lower, and fires were less important at SL than at WOL. Our results suggest that C(4) plants are adapted to warm/dry climatic conditions, but not to extreme droughts, and that the fire regime is controlled by biomass-climate interactions.

Climate↗
PubMed2006

Dual function of an Arabidopsis transcription factor DREB2A in water-stress-responsive and heat-stress-responsive gene expression.

Transcription factor DREB2A interacts with a cis-acting dehydration-responsive element (DRE) sequence and activates expression of downstream genes involved in drought- and salt-stress response in Arabidopsis thaliana. Intact DREB2A expression does not activate downstream genes under normal growth conditions. A negative regulatory domain exists in the central region of DREB2A, and deletion of this region transforms DREB2A to a constitutive active form (DREB2A CA). We carried out microarray analysis of transgenic Arabidopsis-overexpressing DREB2A CA and found that the overexpression of DREB2A CA induces not only drought- and salt-responsive genes but also heat-shock (HS)-related genes. Moreover, we found that transient induction of the DREB2A occurs rapidly by HS stress, and that the sGFP-DREB2A protein accumulates in nuclei of HS-stressed cells. DREB2A up-regulated genes were classified into three groups based on their expression patterns: genes induced by HS, genes induced by drought stress, and genes induced by both HS and drought stress. DREB2A up-regulated genes were down-regulated in DREB2A knockout mutants under stress conditions. Thermotolerance was significantly increased in plants overexpressing DREB2A CA and decreased in DREB2A knockout plants. Collectively, these results indicate that DREB2A functions in both water and HS-stress responses.

Arabidopsis↗
PubMed1999

Differential gene expressions of rice superoxide dismutase isoforms to oxidative and environmental stresses.

Active oxygen species (AOSs) are produced under stress conditions of plant cells. Superoxide dismutase (SOD) catalyzes the first step in the AOS scavenging system. The responses of SOD genes to environmental stresses were analyzed in rice seedlings by the treatments of drought, salinity and chilling. The expressions of abscisic acid (ABA)-inducible genes, Mn-SOD gene (sodA1) and one of the cytosolic Cu/Zn-SOD genes (sodCc2), were strongly induced by the treatment of drought and salinity. While Fe-SOD gene (sodB) and the other cytosolic Cu/Zn-SOD gene (sodCc1) were also induced by ABA. However the mRNA level of sodB was decreased by drought treatment, and sodCc1 gene was not induced by drought and salinity treatments. Plastidic Cu/Zn-SOD gene (sodCp) quickly responded to salinity treatment in the light but not in the dark. In the treatment with hydrogen peroxide, sodCp gene was strongly induced shortly after the treatment. These results suggested that phytohormone and AOSs are associated with the regulation of SOD genes under environmental stresses.

Abscisic Acid↗
PubMed2025

Smarter stomata: emergent technologies unlocking yield potential in a changing climate.

Stomata, the gatekeepers of leaf gas exchange, regulate carbon dioxide uptake and water loss, functions increasingly critical as crops face more frequent, intense heat and drought. Under dry conditions, stomatal conductance (g s) typically decreases, limiting carbon assimilation and yield. Heat stress, in contrast, elicits variable g S responses: sometimes increasing to facilitate transpirational cooling, while at other times decreasing, especially when combined with drought. Heat and drought also induce complex, context-dependent shifts in stomatal anatomy. Smaller, denser stomata improve drought resilience in some cases, while reduced density confers greater tolerance in others. The optimal stomatal ideotype remains unknown, and different or even opposing traits may confer resilience dependent on the environmental scenario. Substantial genotypic variation in g s and stomatal anatomy, high heritability and co-localized quantitative trait loci for stomatal traits and yield highlight their untapped potential as breeding targets for climate-resilient crops. However, stomatal traits remain largely absent from breeding pipelines due to challenges of phenotyping at scale. This is changing rapidly. Advances in deep learning, porometry, digital microscopy, and remote sensing now enable high-throughput measurement of stomatal physiology and anatomy. Next-generation breeding technologies including clustered regularly interspaced short palindromic repeats (CRISPR), multi-omics approaches, and artificial intelligence-driven ideotype selection models could revolutionize breeding, allowing precise engineering of stomatal traits for resilience to environmental stress. The time has come to move beyond characterizing stomatal traits and start actively incorporating them into breeding strategies. By leveraging these technologies, stomatal traits can become high value targets, unlocking their potential to enhance crop performance in a hotter, drier future.

abiotic stress↗
PubMed2003

Interactive effects of water stress and xylem-limited bacterial infection on the water relations of a host vine.

Xylella fastidiosa, a xylem-limited bacterial pathogen that causes bacterial leaf scorch in its hosts, has a diverse and extensive host range among plant species worldwide. Previous work has shown that water stress enhances leaf scorch symptom severity and progression along the stem in Parthenocissus quinquefolia infected by X. fastidiosa. The objective here was to investigate the mechanisms underlying the interaction of water stress and infection by X. fastidiosa. Using the eastern deciduous forest vine, P. quinquefolia, infection and water availability were manipulated while measuring leaf water potentials (psi(L)), stomatal conductance (g(s)), whole shoot hydraulic conductance (K(h)), per cent xylem embolism, and xylem vessel dimensions. No significant differences in any of the physiological measurements were found between control and infected plants prior to drought. Drought treatment significantly reduced psi(L) and g(s) at all leaf positions throughout the day in late summer in both years of the study. In addition, infection significantly reduced psi(L) and g(s) in the most basal leaf positions in late summer in both years. Whole shoot hydraulic conductance was reduced by both low water and infection treatments. However, per cent embolized vessels and mean vessel diameter were affected by drought treatment only. These results imply that the major effect of infection by X. fastidiosa occurs due to reduced hydraulic conductance caused by clogging of the vessels, and not increased cavitation and embolism of xylem elements. The reduced K(h) caused by X. fastidiosa infection acts additively with the water limitation imposed by Drought stress.

Environment↗
PubMed1993

Photosynthesis of oak leaves under water stress: maintenance of high photochemical efficiency of photosystem II and occurrence of non-uniform CO(2) assimilation.

Net CO(2) assimilation rates (A), stomatal conductance to water vapor (g(w)), photosynthetic O(2) evolution in 5% CO(2) (A(max)) and photochemical efficiency of photosystem II were monitored in leaves of young oak saplings (Quercus petraea Matt. Liebl.) in response to increasing drought. Both A and g(w) declined rapidly as soon as predawn leaf water potential dropped below -1.0 MPa. The calculated intercellular concentration of CO(2) first declined and then increased again as drought intensity increased, suggesting that both stomatal closure and a decreased ability of mesophyll chloroplasts to fix available CO(2) were involved in the drought-induced reductions in A and g(w). However, this assumption was not supported by the observations that, with increasing drought, the decline in A(max) was limited, and the photochemistry of photosystem II and the quantum yield of light-driven electron transport remained stable. Autoradiograms of (14)CO(2)-fed leaves revealed non-uniform assimilation rates during water stress. The consequences of a potential artifact induced by this patchiness on the calculation of intercellular CO(2) concentration are discussed.

Journal Article↗
PubMed1994

Adaptive significance of intermittent shoot growth in Douglas-fir seedlings.

We tested the hypothesis that intermittent (lammas) shoot growth in Douglas-fir (Pseudotsuga menziesii var menziesii (Mirb.) Franco) seedlings from dry regions of southwest Oregon is adaptively significant. Seedlings from open-pollinated families (160 total) from two inland (dry) and two coastal (wet) sources were grown under either well-watered or intermittent drought conditions (temporary drought followed by rewatering) for two growing seasons. In the first growing season, the results supported the hypothesis: the frequency of a second flush was genetically controlled (although weakly, h(f) (2) </= 0.34); more seedlings, on average, from inland families than from coastal families displayed a second flush; and seedlings from inland families were more responsive to the intermittent drought regime in terms of increased frequency of a second flush (relative to the frequency in the well-watered regime). During the second growing season, the intermittent drought treatment did not promote intermittent shoot growth, although inland and coastal families had different patterns of shoot growth that reflected adaptations to soil water availability. We conclude that inland families have adapted to dry summers and short growing seasons by relying predominantly on predetermined growth for seedling height increment after the first growing season. In response to wetter and generally longer growing seasons, however, coastal families have developed a less regulated pattern of shoot extension and rely more on free growth.

Journal Article↗
PubMed1996

Isoprene emission, photosynthesis, and growth in sweetgum (Liquidambar styraciflua) seedlings exposed to short- and long-term drying cycles.

Isoprene emissions were studied in one-year old sweetgum (Liquidambar styraciflua L.) seedlings during nine drying-rewatering cycles extending over five months. Each drying cycle lasted to the point of leaf wilting. Growth was essentially stopped in response to the first drying cycle, though seedling survival and capacity to recover turgor on rewatering remained high throughout the entire nine cycles. Photosynthetic rates of leaves were inhibited by the drying treatments. Under severe drought, isoprene emission rates of leaves were also inhibited, though isoprene emission was generally less sensitive to drought than photosynthesis. The lower drought sensitivity of isoprene emission compared with photosynthesis resulted in a higher percentage of fixed carbon lost as isoprene as seedlings became more stressed. During the recovery phase of the drying-rewatering cycles, isoprene emission rates in several seedlings were higher than in well-watered control seedlings. Following the ninth drying-rewatering cycle, sustained daily watering resulted in recovery of isoprene emission rates to control values within four days. Photosynthetic rates only recovered to 50% of control values after seven days. We conclude that the mechanisms regulating photosynthetic rate and isoprene emission rate are differentially influenced by limited water supplies. The results are consistent with past studies that predict a protective role for isoprene emission during stress, particularly protection from excessive leaf temperatures during drought.

Journal Article↗
PubMed1998

Growth, productivity and water use in three hybrid poplar clones.

Three drought treatments (none, medium and severe) were applied to three container-grown, hybrid poplar clones (Beaupré, Trichobel and Ghoy) as a series of drying cycles over two years-in a greenhouse in 1994 and outside in 1995. Leaf area development, annual biomass production and annual water use were recorded. Clone Beaupré had greater rates of leaf area development (up to 11.8 x 10(-4) m(2) degrees C day(-1)) and subsequently greater dry matter production (up to 816 g tree(-1) year(-1)) and water use (up to 168 l tree(-1) year(-1)) than the other two clones. In all clones, rates of leaf area development, annual biomass production and water use were less under drought conditions than under well-watered conditions. On a proportional basis, the highest reduction in biomass by drought was in Trichobel (up to 74%), whereas, in absolute terms, the reduction was greatest in Beaupré (up to 500 g tree(-1)). However, under drought conditions, annual biomass production was greater in Clone Beaupré than in the other clones. The relationship between water use and biomass production was constant over both years, and was similar for Beaupré and Trichobel at 4.4 g l(-1), but was less for Ghoy at 3.5 g l(-1).

Journal Article↗
PubMed1999

Regulation of water loss in populations of Populus trichocarpa: the role of stomatal control in preventing xylem cavitation.

Variations in resistance to drought-induced xylem cavitation, xylem air-entry points, stomatal behavior, and hydraulic conductivity were measured in four populations of Populus trichocarpa Torr. & A. Gray collected along an east-west humidity and temperature gradient in Washington State, USA. Xylem air-entry points were less negative in trees from moist environments (-0.71 and -1.32 MPa in the Hoh and Nisqually populations, respectively) than in trees from dry environments (-1.55 and -1.67 MPa in the Palouse and Yakima populations, respectively). Xylem cavitation in response to experimental drought was consistent with air-injection measures of xylem air-entry points for a given population. Populations vulnerable to cavitation also exhibited higher stem specific hydraulic conductivities and limited stomatal control compared with resistant populations. Populations exhibiting vulnerability to cavitation and limited stomatal control desiccated more rapidly during drought compared with resistant populations. This study provides evidence of interpopulation variation in resistance to drought-induced xylem cavitation, stomatal behavior, and hydraulic conductivity within Populus trichocarpa.

Journal Article↗
PubMed2000

Seasonal CO(2) assimilation and stomatal limitations in a Pinus taeda canopy.

Net CO(2) assimilation (A(net)) of canopy leaves is the principal process governing carbon storage from the atmosphere in forests, but it has rarely been measured over multiple seasons and multiple years. I measured midday A(net) in the upper canopy of maturing loblolly pine (Pinus taeda L.) trees in the piedmont region of the southeastern USA on 146 sunny days over 36 months. Concurrent data for leaf conductance and photosynthetic CO(2) response curves (A(net)-C(i) curves) were used to estimate the relative importance of stomatal limitations to CO(2) assimilation in the field. In fully expanded current-year and 1-year-old needles, midday light-saturated A(net) was constant over much of the growing season (5-6 &mgr;mol CO(2) m(-2) s(-1)), except during drought periods. During the winter season (November-March), midday A(net) of overwintering needles varied in proportion to leaf temperature. Net CO(2) assimilation at light saturation occurred when daytime air temperatures exceeded 5-6 degrees C, as happened on more than 90% of the sunny winter days. In both age classes of foliage, winter carbon assimilation accounted for approximately 15% of the daily carbon assimilation on sunny days throughout the year, and was relatively insensitive to year-to-year differences in temperature during this season. However, strong stomatal limitations to A(net) occurred as a result of water stress associated with freezing cycles in winter. During the growing season, drought-induced water stress produced the largest year-to-year differences in seasonal CO(2) assimilation on sunny days. Seasonal A(net) was more drought sensitive in current-year needles than in 1-year-old needles. Relative stomatal limitations to daily integrated A(net) were approximately 40% over the growing season, and summer drought rather than high temperatures had the largest impact on summer A(net) and integrated annual CO(2) uptake in the upper crown. Despite significant stomatal limitations, a long duration of near-peak A(net) in the upper crown, particularly in 1-year-old needles, conferred high seasonal and annual carbon gain.

Journal Article↗
PubMed2002

Photosynthetic responses to water deficit in six Mediterranean sclerophyll species: possible factors explaining the declining distribution of Rhamnus ludovici-salvatoris, an endemic Balearic species.

We sought to explain the declining distribution in the Balearic Islands of the endemic shrub Rhamnus ludovici-salvatoris R. Chodat, by comparing its photosynthetic response to drought with that of several widely distributed, competing Mediterranean species (R. alaternus L., Quercus ilex L., Pistacia lentiscus L., Q. humilis Mill. and P. terebinthus L.). All of the study species, except for the two Rhamnus species, avoided desiccation by rapidly adjusting their stomatal conductance at the onset of drought, and maintaining constant leaf relative water content. The two Rhamnus species showed desiccation-tolerant behavior; i.e., as drought progressed, their predawn leaf relative water content decreased simultaneously with stomatal closure. All four desiccation-avoiding species showed a significant positive correlation between leaf thermal dissipation (estimated by the fluorescence parameter NPQ (non-photochemical quenching)) and the de-epoxidation state of the xanthophyll cycle (DPS). The two Rhamnus species exhibited maximum DPS regardless of treatment, but only R. alaternus increased NPQ in response to drought. Rhamnus ludovici-salvatoris had a high ratio of photorespiration to photosynthesis and a low intrinsic water-use efficiency; traits that are likely to be unfavorable for plant productivity under arid conditions. It also had the lowest DPS and thermal dissipation among the six species. We conclude that the photosynthetic traits of R. ludovici-salvatoris account for its limited ability to compete with other species in the Mediterranean region.

Carbon Dioxide↗
PubMed2002

Solute accumulation of chestnut oak and dogwood leaves in response to throughfall manipulation of an upland oak forest.

To determine the biochemical basis of osmotic adjustment, seasonal and treatment differences in foliar water- soluble organic solutes and inorganic ions were investigated for two hardwood species that exhibited osmotic adjustment in a Throughfall Displacement Experiment at the Walker Branch Watershed near Oak Ridge, Tennessee. Leaf samples of overstory and understory chestnut oak (Quercus prinus L.) and understory dogwood (Cornus florida L.) were collected during the 1994 (wet) and 1995 (dry) growing seasons from each of three treatments: dry (-33% throughfall), ambient (control) and wet (+33% throughfall). Foliar soluble carbohydrates and organic acids were measured by gas chromatography-mass spectrometry. During May 1994, when the demand for sucrose was greatest, dogwood accumulated small amounts of glucose, quinic acid and Mg2+, offsetting a decline in nitrate concentration. As the mild drought continued and tree growth slowed, there was a significant accumulation of sucrose in dogwood in the dry treatment in June, and a trend toward increased K+. In overstory chestnut oak in the dry treatment over the same period, there were significant accumulations of fructose, glucose and K+, and a trend toward increased quinic acid accumulation. Sucrose did not become a key osmotically active compound in chestnut oak until August 1994. In 1995, with the exception of understory chestnut oak, there was no accumulation of K+ in either species. During the severe drought of 1995, monosaccharides, particularly glucose and fructose, accounted for most of the osmotic adjustment in both species. Among solutes, glucose constituted the largest accumulation in dogwood in the dry treatment in August 1995, followed by fructose and sucrose. There was only a moderate increase in solutes in chestnut oak trees in 1995, with fructose and glucose constituting over 50% of the predicted solute accumulation in July. Both species accumulated a wider array of solutes during the dry year than during the wet year, but treatment differences in solute accumulation in chestnut oak were partially limited by drought. The greater dehydration tolerance of chestnut oak than dogwood was evident in the higher baseline concentrations of organic solutes and the greater array of solutes accumulated in response to drought.

Adaptation, Physiological↗
PubMed2005

Interspecific variation in xylem vulnerability to cavitation among tropical tree and shrub species.

In tropical moist forests, seasonal drought limits plant survival, productivity and diversity. Drought-tolerance mechanisms of tropical species should reflect the maximum seasonal water deficits experienced in a particular habitat. We investigated stem xylem vulnerability to cavitation in nine tropical species with different life histories and habitat associations. Stem xylem vulnerability was scored as the xylem water potential causing 50 and 75% loss of hydraulic conductivity (P50 and P75, respectively). Four shade-tolerant shrubs ranged from moderately resistant (P50=-1.9 MPa for Ouratea lucens Kunth. Engl.) to highly resistant to cavitation (P50=-4.1 MPa for Psychotria horizontalis Sw.), with shallow-rooted species being the most resistant. Among the tree species, those characteristic of waterlogged soils, Carapa guianensis Aubl., Prioria copaifera Griseb. and Ficus citrifolia Mill., were the most vulnerable to cavitation (P50=-0.8 to -1.6 MPa). The wet-season, deciduous tree, Cordia alliodora (Ruiz and Pav.) Oken., had resistant xylem (P50=-3.2 MPa), whereas the dry-season, deciduous tree, Bursera simaruba (L.) Sarg. was among the most vulnerable to cavitation (P50=-0.8 MPa) of the species studied. For eight out of the nine study species, previously reported minimum seasonal leaf water potentials measured in the field during periods of drought correlated with our P50 and P75 values. Rooting depth, deciduousness, soil type and growth habit might also contribute to desiccation tolerance. Our results support the functional dependence of drought tolerance on xylem resistance to cavitation.

Seasons↗
PubMed2006

Water stress-responsive genes in loblolly pine (Pinus taeda) roots identified by analyses of expressed sequence tag libraries.

Drought stress is the principal cause of seedling mortality in pine forests of the southeastern United States and in many other forested regions around the globe. As part of a larger effort to discover loblolly pine genes, this study subjected rooted cuttings of three unrelated pine genotypes to three watering regimens. Expressed sequence tags (ESTs) were obtained from both the 3' and 5' ends of 12,918 randomly selected cDNAs generated from root tissues. These ESTs were clustered to identify 6,765 unique transcripts (UniScripts) derived from 6,202 putative unique genes (UniGenes-S). Tentative annotations were assigned on the basis of BLASTX comparisons to the Protein Information Resource Nonredundant Reference (PIR-NREF) database. Expression levels of 42 UniScripts varied with high statistical significance with respect to treatment. Many of them resembled gene products shown to be important for drought tolerance in other species, including dehydrins, endochitinases, cytochrome P450 enzymes, pathogenesis-related proteins and various late-embryogenesis abundant (LEA) gene products. Similarly, expression levels of 110 UniScripts varied with high statistical significance among genotypes, indicating that gene expression patterns in this species are much more dependent on genotype than on treatment. Most of the water stress-induced pine UniScripts that appeared to encode products resembling drought tolerance factors in other species were most highly induced in a single genotype, suggesting that particularly useful adaptive alleles for drought tolerance might exist within the collection of cDNAs characterized from this genotype. Mining and visualizing the complete data set, as well as downloading of both EST and UniScript contig sequences, are possible using MAGIC Gene Discovery at http://fungen.org/genediscovery/.

Acclimatization↗
PubMed2006

Restoration thinning and influence of tree size and leaf area to sapwood area ratio on water relations of Pinus ponderosa.

Ponderosa pine (Pinus ponderosa Dougl. ex P. Laws) forest stand density has increased significantly over the last century (Covington et al. 1997). To understand the effect of increased intraspecific competition, tree size (height and diameter at breast height (DBH)) and leaf area to sapwood area ratio (A(L):A(S)) on water relations, we compared hydraulic conductance from soil to leaf (kl) and transpiration per unit leaf area (Q(L)) of ponderosa pine trees in an unthinned plot to trees in a thinned plot in the first and second years after thinning in a dense Arizona forest. We calculated kl and Q(L) based on whole- tree sap flux measured with heat dissipation sensors. Thinning increased tree predawn water potential within two weeks of treatment. Effects of thinning on kl and Q(L) depended on DBH, A(L):A(S) and drought severity. During severe drought in the first growing season after thinning, kl and Q(L) of trees with low A(L):A(S) (160-250 mm DBH; 9-11 m height) were lower in the thinned plot than the unthinned plot, suggesting a reduction in stomatal conductance (g(s)) or reduced sapwood specific conductivity (K(S)), or both, in response to thinning. In contrast kl and Q(L) were similar in the thinned plot and unthinned plot for trees with high A(L):A(S) (260-360 mm DBH; 13-16 m height). During non-drought periods, kl and Q(L) were greater in the thinned plot than in the unthinned plot for all but the largest trees. Contrary to previous studies of ponderosa pine, A(L):A(S) was positively correlated with tree height and DBH. Furthermore, kl and Q(L) showed a weak negative correlation with tree height and a strong negative correlation with A(S) and thus A(L):A(S) in both the thinned and unthinned plots, suggesting that trees with high A(L):A(S) had lower g(s). Our results highlight the important influence of stand competitive environment on tree-size-related variation in A(L):A(S) and the roles of A(L):A(S) and drought on whole-tree water relations in response to thinning.

Arizona↗
PubMed1999

Primary forest dynamics in lowland dipterocarp forest at Danum Valley, Sabah, Malaysia, and the role of the understorey.

Changes in species composition in two 4-ha plots of lowland dipterocarp rainforest at Danum, Sabah, were measured over ten years (1986-1996) for trees > or = 10 cm girth at breast height (gbh). Each included a lower-slope to ridge gradient. The period lay between two drought events of moderate intensity but the forest showed no large lasting responses, suggesting that its species were well adapted to this regime. Mortality and recruitment rates were not unusual in global or regional comparisons. The forest continued to aggrade from its relatively (for Sabah) low basal area in 1986 and, together with the very open upper canopy structure and an abundance of lianas, this suggests a forest in a late stage of recovery from a major disturbance, yet one continually affected by smaller recent setbacks. Mortality and recruitment rates were not related to population size in 1986, but across subplots recruitment was positively correlated with the density and basal area of small trees (10-< 50cm gbh) forming the dense understorey. Neither rate was related to topography. While species with larger mean gbh had greater relative growth rates (rgr) than smaller ones, subplot mean recruitment rates were correlated with rgr among small trees. Separating understorey species (typically the Euphorbiaceae) from the overstorey (Dipterocarpaceae) showed marked differences in change in mortality with increasing gbh: in the former it increased, in the latter it decreased. Forest processes are centred on this understorey quasi-stratum. The two replicate plots showed a high correspondence in the mortality, recruitment, population changes and growth rates of small trees for the 49 most abundant species in common to both. Overstorey species had higher rgrs than understorey ones, but both showed considerable ranges in mortality and recruitment rates. The supposed trade-off in traits, viz slower rgr, shade tolerance and lower population turnover in the understorey group versus faster potential growth rate, high light responsiveness and high turnover in the overstorey group, was only partly met, as some understorey species were also very dynamic. The forest at Danum, under such a disturbance-recovery regime, can be viewed as having a dynamic equilibrium in functional and structural terms. A second trade-off in shade-tolerance versus drought-tolerance is suggested for among the understorey species. A two-storey (or vertical component) model is proposed where the understorcy-overstorey species' ratio of small stems (currently 2:1) is maintained by a major feedback process. The understorey appears to be an important part of this forest, giving resilience against drought and protecting the overstorey saplings in the long term. This view could be valuable for understanding forest responses to climate change where drought frequency in Borneo is predicted to intensify in the coming decades.

Ecosystem↗
PubMed2004

Global climate and the distribution of plant biomes.

Biomes are areas of vegetation that are characterized by the same life-form. Traditional definitions of biomes have also included either geographical or climatic descriptors. This approach describes a wide range of biomes that can be correlated with characteristic climatic conditions, or climatic envelopes. The application of remote sensing technology to the frequent observation of biomes has led to a move away from the often subjective definition of biomes to one that is objective. Carefully characterized observations of life-form, by satellite, have been used to reconsider biome classification and their climatic envelopes. Five major tree biomes can be recognized by satellites based on leaf longevity and morphology: needleleaf evergreen, broadleaf evergreen, needleleaf deciduous, broadleaf cold deciduous and broadleaf drought deciduous. Observations indicate that broadleaf drought deciduous vegetation grades substantially into broadleaf evergreen vegetation. The needleleaf deciduous biome occurs in the world's coldest climates, where summer drought and therefore a drought deciduous biome are absent. Traditional biome definitions are quite static, implying no change in their life-form composition with time, within their particular climatic envelopes. However, this is not the case where there has been global ingress of grasslands and croplands into forested vegetation. The global spread of grasses, a new super-biome, was probably initiated 30-45 Myr ago by an increase in global aridity, and was driven by the natural spread of the disturbances of fire and animal grazing. These disturbances have been further extended over the Holocene era by human activities that have increased the land areas available for domestic animal grazing and for growing crops. The current situation is that grasses now occur in most, if not all biomes, and in many areas they dominate and define the biome. Croplands are also increasing, defining a new and relatively recent component to the grassland super-biome. In the case of both grassland and croplands, various forms of disturbance, particularly frequent disturbance, lead to continued range extensions of the biomes.

Climate↗