Allometry of shoot extension units (hereafter termed "current shoots") was analyzed in a Malaysian canopy species, Elateriospermum tapos Bl. (Euphorbiaceae). Changes in current shoot allometry with increasing tree height were related to growth and maintenance of tree crowns. Total biomass, biomass allocation ratio of non-photosynthetic to photosynthetic organs, and wood density of current shoots were unrelated to tree height. However, shoot structure changed with tree height. Compared with short trees, tall trees produced current shoots of the same mass but with thicker and shorter stems. Current shoots with thin and long stems enhanced height growth in short trees, whereas in tall trees, thick and short current shoots may reduce mechanical and hydraulic stresses. Furthermore, compared with short trees, tall trees produced current shoots with more leaves of lower dry mass, smaller area, and smaller specific leaf area (SLA). Short trees adapted to low light flux density by reducing mutual shading with large leaves having a large SLA. In contrast, tall trees reduced mutual shading within a shoot by producing more small leaves in distal than in proximal parts of the shoot stem. The production of a large number of small leaves promoted light penetration into the dense crowns of tall trees. All of these characteristics suggest that the change in current shoot structure with increasing tree height is adaptive in E. tapos, enabling short trees to maximize height growth and tall trees to maximize light capture.
Climate change exposes vegetation to unusual levels of drought, risking a decline in productivity and an increase in mortality. It still remains unclear how trees and forests respond to such unusual drought, particularly Southeast Asian tropical rain forests. To understand leaf ecophysiological responses of tropical rain forest trees to soil drying, a rainfall exclusion experiment was conducted on mature canopy trees of Dryobalanops aromatica Gaertn.f. (Dipterocarpaceae) for 4 months in an aseasonal tropical rain forest in Sarawak, Malaysia. The rainfall was intercepted by using a soft vinyl chloride sheet. We compared the three control and three treatment trees with respect to leaf water use at the top of the crown, including stomatal conductance (gsmax), photosynthesis (Amax), leaf water potential (predawn: Ψpre; midday: Ψmid), leaf water potential at turgor loss point (πtlp), osmotic potential at full turgor (π100) and a bulk modulus of elasticity (ε). Measurements were taken using tree-tower and canopy-crane systems. During the experiment, the treatment trees suffered drought stress without evidence of canopy dieback in comparison with the control trees; e.g., Ψpre and Ψmid decreased with soil drying. Minimum values of Ψmid in the treatment trees decreased during the experiment, and were lower than πtlp in the control trees. However, the treatment trees also decreased their πtlp by osmotic adjustment, and the values were lower than the minimum values of their Ψmid. In addition, the treatment trees maintained gs and Amax especially in the morning, though at midday, values decreased to half those of the control trees. Decreasing leaf water potential by osmotic adjustment to maintain gs and Amax under soil drying in treatment trees was considered to represent anisohydric behavior. These results suggest that D. aromatica may have high leaf adaptability to drought by regulating leaf water consumption and maintaining turgor pressure to improve its leaf water relations.
Variations in leaf photosynthetic, morphological and biochemical properties with increasing plant height from seedlings to emergent trees were investigated in five dipterocarp species in a Malaysian tropical rain forest. Canopy openness increased significantly with tree height. Photosynthetic properties, such as photosynthetic capacity at light saturation, light compensation point, maximum rate of carboxylation and maximum rate of photosynthetic electron transport, all increased significantly with tree height. Leaf morphological and biochemical traits, such as leaf mass per area, palisade layer thickness, nitrogen concentration per unit area, chlorophyll concentration per unit dry mass and chlorophyll to nitrogen ratio, also changed significantly with tree height. Leaf properties had simple and significant relationships with tree height, with few intra- and interspecies differences. Our results therefore suggest that the photosynthetic capacity of dipterocarp trees depends on tree height, and that the trees adapt to the light environment by adjusting their leaf morphological and biochemical properties. These results should aid in developing models that can accurately estimate carbon dioxide flux and biomass production in tropical rain forests.
Forest growth is an important factor both economically and ecologically, and it follows a predictable trend with age. Generally, growth accelerates as canopies develop in young forests and declines substantially soon after maximum leaf area is attained. The causes of this decline are multiple and may be linked to age- or size-related processes, or both. Our objective was to determine the relative effects of tree age and tree size on the physiological attributes of two broadleaf species. As age and size are normally coupled during growth, an approach based on grafting techniques to separate the effects of size from those of age was adopted. Genetically identical grafted seedlings were produced from scions taken from trees of four age classes, ranging from 4 to 162 years. We found that leaf-level net photosynthetic rate per unit of leaf mass and some other leaf structural and biochemical characteristics had decreased substantially with increasing size of the donor trees in the field, whereas other gas exchange parameters expressed on a leaf area basis did not. In contrast, these parameters remained almost constant in grafted seedlings, i.e., scions taken from donor trees with different meristematic ages show no age-related trend after they were grafted onto young rootstocks. In general, the results suggested that size-related limitations triggered the declines in photosynthate production and tree growth, whereas less evidence was found to support a role of meristematic age.
In this study, we demonstrated the occurrence of stomatal patchiness and its spatial scale in leaves from various sizes of trees grown in a lowland dipterocarp forest in Peninsular Malaysia. To evaluate the patterns of stomatal behavior, we used three techniques simultaneously to analyze heterobaric or homobaric leaves from five tree species ranging from 0.6 to 31 m in height: (i) diurnal changes in chlorophyll fluorescence imaging, (ii) observation and simulation of leaf gas-exchange rates and (iii) a pressure-infiltration method. Measurements were performed in situ with 1000 or 500 μmol m(-2) s(-1) photosynthetic photon flux density. Diurnal patterns in the spatial distribution of photosynthetic electron transport rate (J) mapped from chlorophyll fluorescence images, a comparison of observed and simulated leaf gas-exchange rates, and the spatial distribution of stomatal apertures obtained from the acid-fuchsin-infiltrated area showed that patchy stomatal closure coupled with severe midday depression of photosynthesis occurred in Neobalanocarpus heimii (King) Ashton, a higher canopy tree with heterobaric leaves due to the higher leaf temperature and vapor pressure deficit. However, subcanopy or understory trees showed uniform stomatal behavior throughout the day, although they also have heterobaric leaves. These results suggest that the occurrence of stomatal patchiness is determined by tree size and/or environmental conditions. The analysis of spatial scale by chlorophyll fluorescence imaging showed that several adjacent anatomical patches (lamina areas bounded by bundle-sheath extensions within the lamina) may co-operate for the distributed patterns of J and stomatal apertures.
Effects of heterogeneity in stomatal behavior on gas-exchange characteristics of leaves from four tree species growing in different climates, including temperate, tropical monsoon and tropical rain forest, were investigated by combining gas-exchange measurements and the pressure-infiltration method. Field observations indicated linear relationships between whole-leaf conductance and the ratio of infiltrated to non-infiltrated leaf area (open stomata area) in Dipterocarpus sublamellatus Foxw. and Neobalanocarpus heimii (King) Ashton in a tropical rain forest in Peninsular Malaysia, whereas the ratio of infiltrated to non-infiltrated area rapidly increased up to the whole-leaf conductance at which the entire leaf was infiltrated in Cinnamomum camphora Sieb. in a temperate evergreen forest in Japan and in Azadirachta indica Juss. in a tropical monsoon area in Thailand. These results strongly suggest small ranges in bell-shaped stomatal conductance distributions in C. camphora and A. indica and bimodal stomatal conductance distributions in D. sublamellatus and N. heimii. The values of normalized maximum carboxylation rate at 25 degrees C (V(cmax25)) derived from gas-exchange measurements were not constant, but decreased with decreasing whole-leaf conductance in D. sublamellatus and N. heimii. A gas-exchange model analysis revealed a linear relationship between whole-leaf conductance and the ratio of infiltrated to non-infiltrated leaf area for bimodal stomatal conductance distributions, whereas for bell-shaped distributions, the relationships were nonlinear. Midday depression of apparent V(cmax25) in these species was mainly caused by bimodal stomatal closure. The bimodal stomatal distribution model could also explain diurnal changes in photosynthetic assimilation and transpiration rates in these species.
Photosynthetic rate, nitrogen concentration and morphological properties of canopy leaves were studied in 18 trees, comprising five dipterocarp species, in a tropical rain forest in Sarawak, Malaysia. Photosynthetic rate at light saturation (Pmax) differed significantly across species, varying from 7 to 18 micro mol m(-2) s(-1). Leaf nitrogen concentration and morphological properties, such as leaf blade and palisade layer thickness, leaf mass per area (LMA) and surface area of mesophyll cells per unit leaf area (Ames/A), also varied significantly across species. Among the relationships with leaf characteristics, Pmax had the strongest correlation with leaf mesophyll parameters, such as palisade cell layer thickness (r2 = 0.76, P < 0.001) and Ames/A (r2 = 0.73, P < 0.001). Leaf nitrogen concentration and Pmax per unit area also had a significant but weaker correlation (r2 = 0.46, P < 0.01), whereas Pmax had no correlation, or only weakly significant correlations, with leaf blade thickness and LMA. Shorea beccariana Burck, which had the highest P(max) of the species studied, also had the thickest palisade layer, with up to five or more layers. We conclude that interspecific variation in photosynthetic capacity in tropical rain forest canopies is influenced more by leaf mesophyll structure than by leaf thickness, LMA or leaf nitrogen concentration.
Land surface temperature is predicted to increase by 0.2 °C per decade due to climate change, although with considerable regional variability, and heatwaves are predicted to increase markedly in the future. These changes will affect where crops can be grown in the future. Understanding the thermal limits of plant physiological functioning and how flexible such limits are is thus important. Here, we report on the measurements of a core foliar thermotolerance trait, T50, defined as the temperature at which the maximum quantum yield (Fv/Fm) of photosystem II declines by 50%, across nine different Malaysian Hevea brasiliensis clones. We explore the relative importance of interclonal versus intraclonal variation in T50 as well as its association with leaf and hydraulic traits. We find very low variation in T50 within individual clones (mean intraclonal coefficient of variation (CoV) of 1.26%) and little variation across clones (interclonal CoV of 2.1%). The interclonal variation in T50 was lower than for all other functional traits considered. The T50 was negatively related to leaf mass per area and leaf dry matter content, but it was not related to hydraulic traits such as embolism resistance (P50) or hydraulic safety margins (HSM50). The range of T50 observed (42.9-46.2 °C) is well above the current maximum air temperatures Tmax,obs (T50 - Tmax,obs >5.8 °C), suggesting that H. brasiliensis is likely thermally safe in this south-east Asian region of Malaysia.
To explore the potential of in vitro rapid regeneration, three varieties (Golpaygan-181, Orumieh-1763, and Gorgan-1601) of sainfoin (Onobrychis viciifolia Scop. syn. Onobrychis sativa L.) were evaluated. For the first time, an encapsulation protocol was established from somatic embryogenic callus in torpedo and cotyledonary stages to create artificial seeds. Callus derived from different concentrations of Kinetin (0-2.0 mg L(-1)) and Indole-3-acetic acid (0-2.0 mg L(-1)) was coated with sodium alginate and subsequently cultured either in Murashige and Skoog (MS) medium or in soil substrate. Adventitious shoots from synthetic beads developed into rooting in full and half strength MS medium supplemented with various concentrations of auxin and cytokinin. Prolonged water conservation of black and red soils (1:1) had the highest rate of survival plantlets in the acclimatization process. Diverse resistance techniques in Onobrychis viciifolia were evaluated when the plants were subjected to water deficiency. Higher frequency of epicuticular waxes was observed in in vivo leaves compared to in vitro leaves. Jagged trichomes nonsecreting glands covered by spines were only observed in the lower leaf side. Ultimately, stomata indices were 0.127 (abaxial), 0.188 (adaxial) in in vivo and 0.121 (abaxial), 0.201 (adaxial) in in vitro leaves.
Tissue culture studies of Celosia cristata were established from various explants and the effects of various hormones on morphogenesis of this species were examined. It was found that complete plant regeneration occurred at highest percentage on MS medium supplemented with 2.0 mg/L NAA and 1.5 mg/L BAP, with the best response showed by shoot explants. In vitro flowering was observed on MS basal medium after six weeks. The occurrence of somaclonal variation and changes in cellular behavior from in vivo and in vitro grown plants were investigated through cytological studies and image analysis. It was observed that Mitotic Index (MI), mean chromosome numbers, and mean nuclear to cell area ratio of in vitro root meristem cells were slightly higher compared to in vivo values. However, in vitro plants produced lower mean cell areas but higher nuclear areas when compared to in vivo plants. Thus, no occurrence of somaclonal variation was detected, and this was supported by morphological features of the in vitro plants.
Plant functional traits regulate ecosystem functions but little is known about how co-occurring gradients of land use and edaphic conditions influence their expression. We test how gradients of logging disturbance and soil properties relate to community-weighted mean traits in logged and old-growth tropical forests in Borneo. We studied 32 physical, chemical and physiological traits from 284 tree species in eight 1 ha plots and measured long-term soil nutrient supplies and plant-available nutrients. Logged plots had greater values for traits that drive carbon capture and growth, whilst old-growth forests had greater values for structural and persistence traits. Although disturbance was the primary driver of trait expression, soil nutrients explained a statistically independent axis of variation linked to leaf size and nutrient concentration. Soil characteristics influenced trait expression via nutrient availability, nutrient pools, and pH. Our finding, that traits have dissimilar responses to land use and soil resource availability, provides robust evidence for the need to consider the abiotic context of logging when predicting plant functional diversity across human-modified tropical forests. The detection of two independent axes was facilitated by the measurement of many more functional traits than have been examined in previous studies.
Rattan is an important climbing palm taxon in Malaysian tropical rain forests. Many rattan species have unique structures directly associated with certain ant species. In this study, four rattan species (Daemonorops lewisiana, Calamus castaneus, Daemonorops geniculata and Korthalsia scortechinii) were inspected and documented in a field survey concerning their relationships with several ant species. We noticed that two rattan species (D. lewisiana and C. castaneus) were more likely to be associated with ants compared to their neighbouring rattan (Plectomia griffithii). However, D. lewisiana and C. castaneus did not directly provide shelters for ant colonies, but possessed unique structures: upward-pointing spines and funnel-shaped leaves, which are equipped to collect more litter than P. griffithii. To test our litter collecting hypothesis, we measured the inclination of spines from the stem. Our results showed the presence of ant colonies in the litter-collecting rattans (D. lewisiana and C. castaneus), which was significantly higher compared to a non-litter-collecting rattan (P. griffithii). We propose a complex and novel type of adaptation (litter-collection and provision of nesting materials) for rattans, which promotes interactions between the rattan and ants through the arrangements of leaves, leaflets, and spines. In return, the rattan may benefit from ants' services, such as protection, nutrient enhancement, and pollination.
Macaranga bancana is considered as a successful pioneer plant species. Usually found in disturbed and open areas, most of the current research focused on its relations with ants. One of the unique feature of the plants is that the seedling leaves are red, resembling and almost matching the background. Using a portable spectrometer, we measured the color reflectance of M. bancana seedlings (less than 20 cm in height). We also measured the leaf litter reflectance, adult M. bancana leaves and also seedlings of several other species found in the vicinity of M. bancana seedlings. The reflectances of M. bancana seedlings are very similar to that of the leaf litter background. We suggest that this cryptic coloration is crucial during the early stages of the plant when it still cannot rely on the protection of ants.
Rattan spines are most often regarded as an identification trait and perhaps as a physical protection structure. In this study, we study the spinescence traits from five different species rattan: Daemonorops lewisiana, Daemonorops geniculata, Calamus castaneus, Plectomia griffithii, and Korthalsia scortechinii. We tested length, width, angle, strength, spine density, cross-section surface, spine color, and leaf trichomes (only for D. lewisiana, C. castaneus and D. geniculata). We also tested whether the spines were capable of deterring small climbing mammals (for Plectomia griffithii and Calamus castaneus) by using a choice selection experiment. Due to a variety of spine traits, we could not categorize whether any species is more or less spinescent than the others. We suggest that spines have a much more significant role than merely as a physical defense and work together with other rattan characteristics. This is also evidenced by our choice selection experiment, in which the spines on a single stem donot deter small climbing mammals. However, this is a work in progress, and we have outlined several alternative methods to be used in future work.
We have characterized an oil palm (Elaeis guineensis Jacq.) constitutive promoter that is derived from a translationally control tumor protein (TCTP) gene. The TCTP promoter was fused transcriptionally with the gusA reporter gene and transferred to monocot and dicot systems in order to study its regulatory role in a transient expression study. It was found that the 5' region of TCTP was capable of driving the gusA expression in all the oil palm tissues tested, including immature embryo, embryogenic callus, embryoid, young leaflet from mature palm, green leaf, mesocarp and stem. It could also be used in dicot systems as it was also capable of driving gusA expression in tobacco leaves. The results indicate that the TCTP promoter could be used for the production of recombinant proteins that require constitutive expression in the plant system.
Photosynthetic acclimation (photoacclimation) is the process whereby leaves alter their morphology and/or biochemistry to optimize photosynthetic efficiency and productivity according to long-term changes in the light environment. The three-dimensional architecture of plant canopies imposes complex light dynamics, but the drivers for photoacclimation in such fluctuating environments are poorly understood. A technique for high-resolution three-dimensional reconstruction was combined with ray tracing to simulate a daily time course of radiation profiles for architecturally contrasting field-grown wheat (Triticum aestivum) canopies. An empirical model of photoacclimation was adapted to predict the optimal distribution of photosynthesis according to the fluctuating light patterns throughout the canopies. While the photoacclimation model output showed good correlation with field-measured gas-exchange data at the top of the canopy, it predicted a lower optimal light-saturated rate of photosynthesis at the base. Leaf Rubisco and protein contents were consistent with the measured optimal light-saturated rate of photosynthesis. We conclude that, although the photosynthetic capacity of leaves is high enough to exploit brief periods of high light within the canopy (particularly toward the base), the frequency and duration of such sunflecks are too small to make acclimation a viable strategy in terms of carbon gain. This suboptimal acclimation renders a large portion of residual photosynthetic capacity unused and reduces photosynthetic nitrogen use efficiency at the canopy level, with further implications for photosynthetic productivity. It is argued that (1) this represents an untapped source of photosynthetic potential and (2) canopy nitrogen could be lowered with no detriment to carbon gain or grain protein content.
KEY MESSAGE: Morphological and transcriptomic evidences provide us strong support for the function of ANAC019 in reproductive development under drought stress. Plants are sensitive to drought conditions, particularly at the reproductive stage. Several studies have reported drought effects on crop reproductive development, but the molecular mechanism underlying drought response during reproduction is still unclear. A recent study showed that drought induces in Arabidopsis inflorescence increased expression of many genes, including ANAC019. However, the function of ANAC019 in drought response during reproductive development has not been characterized. Here, we report an investigation of the ANAC019 function in the response to drought during reproduction. ANAC019 is preferentially expressed in the inflorescence compared with the leaf, suggesting possible roles in regulating both stress response and flower development. The anac019 mutant was more sensitive to drought than WT plant, and exhibited a delay in recovery of floral organ development under prolonged drought stress. Moreover, many fewer genes were differentially expressed in the anac019 inflorescence under drought than that of WT, suggesting that the mutant was impaired in drought-induced gene expression. The genes affected by ANAC019 were associated with stress and hormone responses as well as floral development. In particular, the expression levels of several key drought-induced genes, DREB2A, DREB2B, ARF2, MYB21 and MYB24, were dramatically reduced in the absence of ANAC019, suggesting that ANAC019 is an upstream regulator these genes for drought response and flower development. These results provide strong support for the potential function of ANAC019 in reproductive development under drought stress.
South East Asia is widely regarded as a centre of threatened biodiversity owing to extensive logging and forest conversion to agriculture. In particular, forests degraded by repeated rounds of intensive logging are viewed as having little conservation value and are afforded meagre protection from conversion to oil palm. Here, we determine the biological value of such heavily degraded forests by comparing leaf-litter ant communities in unlogged (natural) and twice-logged forests in Sabah, Borneo. We accounted for impacts of logging on habitat heterogeneity by comparing species richness and composition at four nested spatial scales, and examining how species richness was partitioned across the landscape in each habitat. We found that twice-logged forest had fewer species occurrences, lower species richness at small spatial scales and altered species composition compared with natural forests. However, over 80 per cent of species found in unlogged forest were detected within twice-logged forest. Moreover, greater species turnover among sites in twice-logged forest resulted in identical species richness between habitats at the largest spatial scale. While two intensive logging cycles have negative impacts on ant communities, these degraded forests clearly provide important habitat for numerous species and preventing their conversion to oil palm and other crops should be a conservation priority.
Parental distance and plant density dependence of seedling leaf turnover and survival was examined to investigate predictions of the Janzen-Connell hypothesis. The focal study species, Shorea macroptera is a canopy tree species in a lowland rain forest in peninsular Malaysia. We found that the peak of the distribution of plants shifted from 3-6 m to 6-9 m during the course of the change from seedling to sapling stage. The leaf demography of the seedlings was influenced by their distance from the adult tree and also by the seedling density. Although significant density- and distance dependence in leaf production was not detected, seedling leaf loss decreased with distance from the parent tree and with seedling density. Similarly, leaf damage was not found to be distance- or density-dependent, but net leaf gain of seedlings increased with distance from the parent tree. Although no significant distance- or density-dependence was evident in terms of leaf damage, significant distance dependence of the net leaf gain was found. Thus, we concluded that positive distance dependence in the leaf turnover of seedlings may gradually contribute to a shift in the distribution pattern of the progeny through reductions in growth and survivorship.
Vertical variation in leaf gas exchange characteristics of trees grown in a lowland dipterocarp forest in Peninsular Malaysia was investigated. Maximum net photosynthetic rate, stomatal conductance, and electron transport rate of leaves at the upper canopy, lower canopy, and forest floor were studied in situ with saturated condition photosynthetic photon flux density. The dark respiration rate of leaves at the various heights was also studied. Relationships among gas exchange characteristics, and also with nitrogen content per unit leaf area and leaf dry matter per area were clearly detected, forming general equations representing the vertical profile of several important parameters related to gas exchange. Numerical analysis revealed that the vertical distribution of gas exchange parameters was well determined showing both larger carbon gain for the whole canopy and at the same time positive carbon gain for the leaves of the lowest layer. For correct estimation of gas exchange at both leaf and canopy scales using multi-layer models, it is essential to consider the vertical distribution of gas exchange parameters with proper scaling coefficients.