Z.Q. Cai

Seasonal differences in leaf-level physiology give lianas a competitive advantage over trees in a tropical seasonal forest

Lianas are an important component of most tropical forests, where they vary in abundance from high in seasonal forests to low in aseasonal forests. We tested the hypothesis that the physiological ability of lianas to fix carbon (and thus grow) during seasonal drought may confer a distinct advantage in seasonal tropical forests, which may explain pan-tropical liana distributions. We compared a range of leaf-level physiological attributes of 18 co-occurring liana and 16 tree species during the wet and dry seasons in a tropical seasonal forest in Xishuangbanna, China. We found that, during the wet season, lianas had significantly higher CO2 assimilation per unit mass (Amass), nitrogen concentration (Nmass), and δ13C values, and lower leaf mass per unit area (LMA) than trees, indicating that lianas have higher assimilation rates per unit leaf mass and higher integrated water-use efficiency (WUE), but lower leaf structural investments. Seasonal variation in CO2 assimilation per unit area (Aarea), phosphorus concentration per unit mass (Pmass), and photosynthetic N-use efficiency (PNUE), however, was significantly lower in lianas than in trees. For instance, mean tree Aarea decreased by 30.1% from wet to dry season, compared with only 12.8% for lianas. In contrast, from the wet to dry season mean liana δ13C increased four times more than tree δ13C, with no reduction in PNUE, whereas trees had a significant reduction in PNUE. Lianas had higher Amass than trees throughout the year, regardless of season. Collectively, our findings indicate that lianas fix more carbon and use water and nitrogen more efficiently than trees, particularly during seasonal drought, which may confer a competitive advantage to lianas during the dry season, and thus may explain their high relative abundance in seasonal tropical forests.

Leaf development and photosynthetic properties of three tropical tree species with delayed greening

Leaf developmental patterns were characterized for three tropical tree species with delayed greening. Changes in the pigment contents, photosynthetic capacity, stomata development, photosystem 2 efficiency, rate of energy dissipation, and the activity of partial protective enzymes were followed in developing leaves in an attempt to elucidate the relative importance of various photoprotective mechanisms during leaf ontogeny. Big leaves of Anthocephalus chinensis, a fast-growing light demanding species, expanded following an exponential pattern, while relatively small leaves of two shade-tolerant species Litsea pierrei and Litsea dilleniifolia followed a sigmoidal pattern. The juvenile leaves of A. chinensis and L. pierrei contained anthocyanin located below the upper epidermis, while L. dilleniifolia did not contain anthocyanin. Leaves of A. chinensis required about 12 d for full leaf expansion (FLE) and photosynthetic development was delayed 4 d, while L. pierrei and L. dilleniifolia required 18 or 25 d for FLE and photosynthetic development was delayed 10 or 15 d, respectively. During the leaf development the increase in maximum net photosynthetic rate was significantly related to changes in stomatal conductance and the leaf maturation period was positively related to the steady-state leaf dry mass per area for the three studied species. Dark respiration rate of leaves at developing stages was greater, and pre-dawn initial photochemical efficiency was lower than that of mature leaves. Young leaves displayed greater energy dissipation than mature leaves, but nevertheless, the diurnal photoinhibition of young L. dilleniifolia leaves was higher than that of mature leaves. The young red leaves of A. chinensis and L. pierrei with high anthocyanin contents and similar diurnal photoinhibition contained more protective enzymes (superoxide dismutase, ascorbate peroxidase) than mature leaves. Consequently, red leaves may have higher antioxidant ability.

Contrasting nitrogen and phosphorus resorption efficiencies in trees and lianas from a tropical montane rain forest in Xishuangbanna, south-west China

Tropical montane rain forest is widely considered to be a highly threatened hotspot of global diversity (Brummitt & Nic Lughadha 2003), and one of the least understood humid tropical forest ecosystems in terms of nutrient cycling (Bruijnzeel & Proctor 1995). There is, therefore, an urgent need to improve our understanding of nutrient cycling processes in this ecosystem, including the absorption of nutrients (mainly N and P) from senescing leaves, which may be a key component of adaptive mechanisms that conserve limiting nutrients (Killingbeck 1996). Nutrients which are not resorbed, however, will be circulated through litterfall in the longer term (Aerts 1996). The degree of nutrient resorption affects litter quality, which consequently affects decomposition rates and soil nutrient availability (Aerts & Chapin 2000). The importance of resorption in nutrient conservation has led to general hypotheses that species adapted to nutrient-poor environments have high resorption efficiencies (Richardson et al. 2005), and that low leaf nutrient concentrations are associated with high resorption efficiencies within species (Aerts 1996, Kobe et al. 2005). Nutrient resorption has also been shown not to differ greatly between growth forms (e.g. shrubs, grasses, forbs and trees) (Aerts 1996). However, its relative importance among plant functional groups is still highly controversial (Richardson et al. 2005).

Above- and below-ground competition in high and low irradiance: tree seedling responses to a competing liana Byttneria grandifolia

In tropical forests, trees compete not only with other trees, but also with lianas, which may limit tree growth and regeneration. Liana effects may depend on the availability of above- and below-ground resources and differ between tree species. We conducted a shade house experiment to test the effect of light (4% and 35% full sun, using neutral-density screen) on the competitive interactions between seedlings of one liana (Byttneria grandifolia) and three tree species (two shade-tolerant trees, Litsea dilleniifolia and Pometia tomentosa, and one light-demanding tree, Bauhinia variegata) and to evaluate the contribution of both above- and below-ground competition. Trees were grown in four competition treatments with the liana: no competition, root competition, shoot competition and root and shoot competition. Light strongly affected leaf photosynthetic capacity (light-saturated photosynthetic rate, Pn), growth and most morphological traits of the tree species. Liana-induced competition resulted in reduced Pn, total leaf areas and relative growth rates (RGR) of the three tree species. The relative importance of above- and below-ground competition differed between the two light levels. In low light,RGR of the threetree species was reduced more strongly by shoot competition (23.1-28.7% reduction) than by root competition (5.3-26.4%). In high light, in contrast, root competitionratherthanshootcompetitiongreatlyreducedRGR.Lianacompetitionaffectedmostmorphologicaltraits (except for specific leaf area and leaf area ratio of Litsea and Pometia), and differentially altered patterns of biomass allocation in the tree seedlings. These findings suggest that competition from liana seedlings can greatly suppress growth in tree seedlings of both light-demanding and shade-tolerant species and those effects differ with competition type (below- and above-ground) and with irradiance.

Synergistic effect of colonization with arbuscular mycorrhizal fungi improves growth and drought tolerance of Plukenetia volubilis seedlings

The objective of this study was to investigate the effects of arbuscular mycorrhizal fungus (AMF) inoculation on plant growth and drought tolerance in seedlings of a promising oilseed crop, Sacha Inchi (Plukenetia volubilis L.), under well-watered or drought conditions. AMF inoculation was applied in four treatments: without AMF inoculation, Glomus versiforme, Paraglomus occultum, or combination of both microorganism inoculations. The results showed that AMF colonization significantly enhanced the growth of Sacha Inchi seedlings regardless of soil water conditions, and the greatest development was reached in plants dually inoculated under well-watered conditions. G. versiforme was more efficient than P. occultum. Plants inoculated with both symbionts had significantly greater specific leaf area, leaf area ratio and root volume when compared with the uninoculated control, G. versiforme, and P. occultum treatments alone, indicating a synergistic effect in the two AMF inoculation. Photosynthetic rate and water-use efficiency were stimulated by AMF, but not stomatal conductance. Inoculation with AM fungus increased antioxidant enzymes activities including guaiacol peroxidase and catalase, thus lowering hydrogen peroxide accumulation and oxidative damage, especially under drought stress conditions. However, proline content showed little change during drought stress and AMF colonization conditions, which suggested that proline accumulation might not serve as the main compound for osmotic adjustment of the studied species. These results indicate that AMF inoculation stimulated growth and enhanced drought tolerance of Sacha Inchi seedlings, through alterations in morphological, physiological and biochemical traits. This microbial symbiosis might be an effective cultivation practice in improving the performance and development for Sacha Inchi plants.

Responses of two field-grown coffee species to drought and re-hydration

The gas exchange, parameters of chlorophyll fluorescence, contents of pigments, and activities of superoxide dismutase (SOD), ascorbate peroxidase (APX), as well as lipid peroxidation were investigated in two field-grown coffee species, Coffea arabica and C. liberica, exposed to drought and re-hydration. Drought caused a more pronounced inhibition of net photosynthetic rate in C. liberica compared to C. arabica. The de-epoxidation of xanthophyll cycle pigments at midday estimated by leaf reflectance was much higher in C. arabica than in C. liberica, but no significant change was found in response to drought. Under moderate drought, the activities of SOD and APX increased significantly only in C. arabica. The maximum photochemical efficiency of photosystem 2, PS2 (Fv/Fm) at predawn did not change and there was no lipid peroxidation during this time. Under severe drought Fv/Fm decreased and initial fluorescence (F0) increased for both species, and SOD activity increased, APX activity remained relatively high, and malondialdehyde (MDA) accumulated in C. arabica, while APX decreased in C. liberica. The photosynthetic apparatus of C. arabica was completely recovered after 5 d of re-irrigation as indicated by the restoration of Fv/Fm to the control values. A lack of recovery upon rewatering of C. liberica indicated irreversible damage to PS2. Hence compared to C. liberica, C. arabica possesses a higher desiccation-induced antioxidative protection and higher portion of the total pigment pool used in photoprotection, which might aid alleviating photoinhibitory damage during desiccation and photosynthesis recovery when favourable conditions are restored.

Vegetative growth and photosynthesis in coffee plants under different watering and fertilization managements in Yunnan, SW China

In a field experiment Coffea arabica L. was subjected to various moisture and fertilizer regimes in Simao, Yunan, SW China. The experimental treatments consisted of eight factorial combinations of two fertilization levels (high and low) and four watering treatments applied in the dry season: application of dry rice straw mulch, drip irrigation, mulching plus drip irrigation on the soil surface, and control (no mulching or irrigation). The growth of the coffee plants was monitored throughout the course of a full year. Two clear growth peaks were detected (one at the beginning and one in the middle of the wet season) in plants subjected to all treatments, and the growth rhythm of coffee plants was not regulated by extrinsic abiotic factors. High fertilization resulted in a significantly higher relative growth rates for both height and length of the branches during the growth peaks than the low fertilization treatment. In the dry season, increasing the soil moisture contents by irrigation and/or mulching enhanced the plants’ gas exchange, but the soil water status had no significant effects on the internal fluorescence parameters of photosystem 2. More fertilized plants had a greater ability to acclimate to high-irradiance environments than the lightly fertilized plants, showing significant lower diurnal photoinhibition, associated with higher energy utilization through photochemistry and energy dissipation through the xanthophyll cycle. Hence the wet season is the optimum period for photosynthetic carbon fixation and vegetative growth of coffee plants. Higher than routinely applied levels of fertilization are required to optimize the coffee plants’ photosynthetic acclimation and growth in the studied environment. Both soil moisture conserving practices tested, mulching and drip irrigation, had significant effects on the growth and photosynthesis of the coffee plants, but the former was more practical than the latter.

Contrasting leaf characteristics of trees and lianas in secondary and mature forests in southwestern China

We compared variation in sun-canopy leaf anatomy, morphology and photosynthetic rates of coexisting woody species (trees and lianas) in an 8-year-old secondary forest (SF) and mature forest (MF) in the wet season in Xishuangbanna, SW China. Variability of leaf traits of 66 species within growth-form groups in each forest was quantified using coefficients of variation (CV). For the mean values, the woody species in the SF had significantly higher leaf thickness and stomatal density, but lower nonmesophyll/mesophyll ratios than those in the MF. The average leaf area and leaf mass area (LMA) in the studied woody species did not change greatly during the successional process, but differed significantly between the growth forms, with trees having higher values than lianas. The light-saturated photosynthetic rate per unit leaf area (Aa) of the woody species in the SF ranged from 11.2 to 34.5 μmol m−2 s−1, similarly to pioneer tree species from literature data in southeast Asia. The Aa and photosynthetic nitrogen-use efficiency (PNUE) were significantly higher than those in the MF; whereas Aa in the MF ranged between 9 to 21 μmol m−2 s−1, with similar values between lianas and trees. For all woody species in both SF and MF, there were no significant differences in the average values of the CV of all measured variables for both lianas and trees. However, considerable variation in leaf anatomy, morphology, and photosynthetic rates within both growth forms and forests existed, as well as substantial variation in leaf size and stomatal density. We concluded that the tropical woody species formed a heterogeneous functional group in terms of leaf morphology and physiology in both secondary and mature forests.

Chromosome number variation in a promising oilseed woody crop, Plukenetia volubilis L. (Euphorbiaceae)

Cytology of a promising oilseed woody crop, Plukenetia volubilis, was analyzed for the first time here. There are in total 11 different chromosome numbers, ranging from 2n = 50 to 2n = 86 in P. volubilis. Different numbers were found not only in different root tips but also in different cells of the same tip. Among the cells, 2.78% cells had the lowest chromosome number of 2n = 50; and 1.39% cells had the highest chromosome number of 2n = 86. The most common chromosome number is 2n = 58 and about 27.79% cells had this quantity of chromosomes. The chromosomes were extremely small in size, at less than 2 μm. Thus, analysis of chromosome morphology was impaired. Polyploidy, confirmed as the significant evolutionary trend in chromosome number within this species, is briefly discussed. This species is a problematic and polymorphic taxon and our results indicate that its polyploidy can be regarded as the reason.

Planting density and fertilisation independently affect seed and oil yields in Plukenetia volubilis L. plants

Summary To investigate the effects of planting density and fertiliser level on some important agronomic traits of Plukenetia volubilis, a perennial oilseed species, a field experiment with four replications in a completely randomised block design was conducted in a tropical region of China in the 2012-2013 growing season. Planting density (1,666, 2,500, 4,444, 5,000, or 10,000 plants ha–1) was assigned to the main plots and a 1:1:1 (w/w/w) NPK fertiliser at 0, 50 100, 150, or 200 kg ha–1 was assigned factorially to the sub-plots. The results showed that neither planting density nor the rate of fertilisation affected seed size (dry matter) and the phenological development of P. volubilis plants, including the times of initial flowering and maturity, or the pattern of fruit ripening. Planting density, rather than fertiliser level, significantly affected seed oil content, but a non-linear effect was found. Across the different treatments and sampling times, seed size and oil content had relatively high and constant heritabilities indicated by low coefficients of variation. Total seed yields and total oil yields per plot over the growing season ranged from 1,340 - 2,486 kg ha–1, and from 501 - 899 kg ha–1, respectively, with a quadratic response to planting density at all fertiliser levels. Total seed and oil yields increased continuously with the increasing levels of fertiliser used.The absence of any planting density fertiliser level interaction on total seed and oil yields suggested that planting density and fertilisation had independent effects, and that seed or oil yield responses to planting density were not significantly affected by fertiliser level. We concluded that oil production in P. volubilis plants required high levels of fertiliser and, regardless of fertiliser level, a planting density of approx. 4,444 plants ha–1 was required to ensure maximum yield in the field.