Undamaged cotton plants yield more if their neighbour is damaged: implications for pest management.

Understanding the compensatory responses of crops to pest damage is important in developing pest thresholds. Compensation for pest damage in crops can occur at the plant level, where the architecture, growth dynamics and allocation patterns of damaged plants are altered, allowing them to recover or, at the crop level, where differential damage between plants may alter plant-to-plant interactions. We investigated growth and yield of cotton (Gossypium hirsutum L.) following non-uniform manual defoliation of seedlings. This partially replicates real pest damage and is valuable in understanding crop-level responses to damage because it can be inflicted precisely. Damage distributions included damaging 0, 25, 50, 75 or 100% of the plants. Damage intensity for the damaged plants was varied by removing 100 or 75% of each true leaf when plants had two, four and six true leaves. At the crop level, yield loss increased as the proportion of plants damaged and intensity of damage per damaged plant increased. Neighbour interactions occurred; undamaged plants with damaged neighbours grew larger and yielded better than undamaged plants with undamaged neighbours, while the converse applied for damaged plants with undamaged neighbours. Neighbour interactions were influenced by the intensity of damage and were stronger when 100% of the leaf area was removed than when 75% was removed. At the crop level, when compared with yield estimates based on yield of plants from uniformly damaged or undamaged plots, these interactions resulted in higher yield than expected (+8%). This suggests that damage distribution may have to be considered in studies where artificial or real pest damage is inflicted uniformly on plants.

Circumvention of over-excitation of PSII by maintaining electron transport rate in leaves of four cotton genotypes developed under long-term drought.

We investigated the patterns of response to a long-term drought in the field in cotton cultivars (genotypes) with known differences in their drought tolerance. Four cotton genotypes with varying physiological and morphological traits, suited to different cropping conditions, were grown in the field and subjected to a long-term moderate drought. In general, cotton leaves developed under drought had significantly higher area-based leaf nitrogen content (N (area)) than those under well irrigation. Droughted plants showed a lower light-saturated net photosynthetic rate (A (sat)) with lower stomatal conductance (g (s)) and intercellular CO (2) concentration (C (i)) than irrigated ones. Based on the responses of A (sat) to g (s) and C (i), there was no decreasing trend in A (sat) at a given g (s) and C (i) in droughted leaves, suggesting that the decline in A (sat) in field-grown cotton plants under a long-term drought can be attributed mainly to stomatal closure, but not to nonstomatal limitations. There was little evidence of an increase in thermal energy dissipation as indicated by the lack of a decrease in the photochemical efficiency of open PSII (F (v)'/F (m)') in droughted plants. On the basis of electron transport (ETR) and photochemical quenching (q (P)), however, we found evidence indicating that droughted cotton plants can circumvent the risk of excessive excitation energy in photosystem (PS) II by maintaining higher electron transport rates associated with higher N (area), even while photosynthetic rates were reduced by stomatal closure.

Leaf morphology and photosynthetic adjustments among deciduous broad-leaved trees within the vertical canopy profile.

Photosynthetic acclimation of deciduous broad-leaved tree species was studied along a vertical gradient within the canopy of a multi-species deciduous forest in northern Japan. We investigated variations in (1) local light regime and CO2 concentration ([CO2]), and (2) morphological (area, thickness and area per mass), biochemical (nitrogen and chlorophyll concentrations) and physiological (light-saturated photosynthetic rate) attributes of leaves of seven major species on three occasions (June, August and October). We studied early successional species, alder (Alnus hirsuta (Spach) Rupr.) and birch (Betula platyphylla var. japonica (Miq.) Hara); gap phase species, walnut (Juglans ailanthifolia Carrière) and ash (Fraxinus mandshurica var. japonica Rupr.); mid-successional species, basswood (Tilia japonica (Miq.) Simonk.) and elm (Ulmus davidiana var. japonica (Rehd.) Nakai); and the late-successional species, maple (Acer mono Bunge). All but maple initiated leaf unfolding from the lower part of the crown. The [CO2] within the vertical profile ranged from 320-350 ppm in the upper canopy to 405-560 ppm near the ground. The lowest and highest ambient [CO2] occurred during the day and during the night, respectively. This trend was observed consistently during the summer, but not when trees were leafless. Chlorophyll concentration was positively related to maximum photosynthetic rate within, but not among, species. Leaf senescence started from the inner part of the crown in alder and birch, but started either in the outer or top portion of the canopy of ash, basswood and maple. Chlorophyll (Chl) to nitrogen ratio in leaves increased with decreasing photon flux density. However, Chl b concentration in all species remained stable until the beginning of leaf senescence. Maximum photosynthetic rates observed in sun leaves of early successional species, gap phase or mid-successional species, and late successional species were 12.5-14.8 micromol m(-2) s(-1), 4.1-7.8 micromol m(-2) s(-1) and 3.1 micromol m(-2) s(-1), respectively.

Manganese toxicity as indicated by visible foliar symptoms of Japanese white birch (Betula platyphylla var. japonica).

For the purpose of a field diagnosis of Mn toxicity, we showed the possibility of using visible foliar symptoms of Japanese white birch (Betula platyphylla var. japonica Hara) as indicator. To examine the relationship between the expression of visible symptoms and leaf Mn concentrations, white birch seedlings were grown under four different Mn levels: 1 mg Mn l-1 as control, 10, 50 and 100 mg Mn l-1. Foliar symptoms of Mn toxicity for white birch were: (1) chlorosis at entire young leaves in the 50 and 100 mg Mn l-1 treatments; and (2) brown speckles at the leaf marginal and interveinal area for old leaves in the treatments greater than 1 mg Mn l-1. Mn preferably accumulated into the leaf marginal and interveinal area, where the brown speckles were observed. The mechanism determining the expression of symptoms seems to be associated with the physiological state related to leaf age as well as Mn distribution and concentration within a leaf.

Inhibition of seedling survival under Rhododendron maximum (Ericaceae): could allelopathy be a cause?

In the southern Appalachian mountains a subcanopy species, Rhododendron maximum, inhibits the establishment and survival of canopy tree seedlings. One of the mechanisms by which seedlings could be inhibited is an allelopathic effect of decomposing litter or leachate from the canopy of R. maximum (R.m.) on seed germination, root elongation, or mycorrhizal colonization. The potential for allelopathy by R.m. was tested with two bioassay species (lettuce and cress), with seeds from four native tree species, and with three ectomycorrhizal fungi. Inhibitory influences of throughfall, fresh litter, and decomposed litter (organic layer) from forest with R.m. (+R.m. sites) were compared to similar extractions made from forest without R.m. (-R.m. sites). Throughfall and leachates of the organic layer from both +R.m. and -R.m. sites stimulated germination of the bioassay species above that of the distilled water control, to a similar extent. There was an inhibitory effect of leachates of litter from +R.m. sites on seed germination and root elongation rate of both bioassay species compared with that of litter from -R.m. sites. Native tree seed stratified in forest floor material from both forest types had a slightly higher seed germination rate compared with the control. A 2-yr study of seed germination and seedling mortality of two tree species, Quercus rubra and Prunus serotina, in field plots showed no significant influence of litter or organic layer from either forest type. Incorporating R.m. leaf material into the growth medium in vitro depressed growth of one ectomycorrhizal species but did not affect two other species. Leaf material from other deciduous tree species depressed ectomycorrhizal growth to a similar or greater extent as leaf material from R.m. In conclusion, R.m. litter can have an allelopathic effect on seed germination and root elongation of bioassay species as well as some ectomycorrhizal species. However, this allelopathic affect is not manifest in field sites and is not likely to be an important cause for the inhibition of seedling survival within thickets of R.m.

Application of chlorophyll fluorescence to evaluate Mn tolerance of deciduous broad-leaved tree seedlings native to northern Japan.

We used chlorophyll fluorescence to examine photosynthetic responses to excess Mn accumulation in leaves of four tree species differing in successional traits. Betula ermanii Cham. (Be) and Alnus hirsuta Turcz. (Ah) were studied as representatives of early-successional species. Ulmus davidiana Planch. var. japonica (Rehder) Nakai (Ud) was selected as a mid-successional species, and Acer mono Maxim. var. glabrum (Lév. et Van't.) Hara (Am) was chosen as a late-successional species. In Be, Ah and Am, high foliar concentrations of Mn had little effect on maximum photochemical efficiency of photosystem II (PSII), as indicated by the values of dark-adapted F(v)/F(m), whereas a significant decrease was observed in Ud. Photochemical quenching (qP) and the excitation capture efficiency of open PSII (F'(v)/F'(m)) decreased with increasing leaf Mn concentration at photosynthetic steady state after a 15-min exposure to 430 &mgr;mol m(-2) s(-1) PPFD. Compared with early-successional species, these decreases were greater in mid- and late-successional species leading to lower effective quantum efficiencies of PSII (DeltaF/F'(m) = qP x F'(v)/F'(m) = (F'(m) - F)/F'(m)). To determine the extent of photoinhibition, F(v)/F(m) of the illuminated leaves was remeasured after a 15-min dark period. Compared with the dark-adapted F(v)/F(m), we observed a significant decrease in F(v)/F(m) in Am leaves containing high concentrations of Mn. These chlorophyll fluorescence studies indicate that the early-successional species Be and Ah have a higher tolerance to excessive accumulations of Mn in leaves than the mid- and late-successional species Ud and Am.

Comparison of photosynthetic responses to manganese toxicity of deciduous broad-leaved trees in northern Japan.

The effects of manganese (Mn) toxicity on photosynthesis of four tree species in northern Japan representing different successional traits were examined. The four species are: Betula ermanii (Be) and Alnus hirsuta (Ah) representing two early successional species, Ulmus davidiana var. japonica (Ud) as the mid-successional species, and Acer mono (Am) as the late successional species. Seedlings were grown hydroponically in a solution containing nutrients and Mn of four concentrations (1, 10, 50, 100 mg litre(-1)) for 50 days. Gas exchange measurements indicate that in all species, Mn accumulation in leaves resulted in the decline of light-saturated net photosynthetic rate ai ambient CO(2) pressure (35 Pa, Pn(amb)) and at saturating (5%) CO(2) pressure (Pn(sat)), and of carboxylation efficiency but has little effect on the maximum efficiency of photochemistry. Sensitivity to elevated levels of Mn differed among species where the decline of Pn(amb) was much more modest in the two early successional species of Be and Ah than the mid- and late successional species of Ud and Am. The same trends were observed in both Pn(sat) and carboxylation efficiency. Based on these results, we suggest that early successional species (Betula ermanii and Alnus hirsuta) have greater tolerance for excess Mn in leaves than mid- and late successional species.

Comparison of the photosynthetic capacity of Siberian and Japanese birch seedlings grown in elevated CO(2) and temperature.

The effects of increased CO(2) and temperature on the photosynthetic capacity of Siberian white birch and Japanese white birch (Betula platyphylla Sukatch. and B. platyphylla Sukatch. var. japonica Hara) were measured. Birch seedlings were raised with a CO(2) partial pressure of 36 +/- 0.3 Pa (i.e., ambient) or 70 +/- 0.6 Pa at day/night temperatures of either 30/16 degrees C or 26/12 degrees C. Siberian birch leaves were smaller and thicker than Japanese birch leaves. Water use efficiency and nitrogen use efficiency of Siberian birch grown in the CO(2)-enriched air were higher than those of Japanese birch. Both species showed a physiological adjustment to the growth CO(2) partial pressure. Carboxylation efficiency and quantum yield of both species grown in CO(2)-enriched air were lower than those of seedlings grown in ambient CO(2). The adaptation of Siberian and Japanese birch to elevated CO(2) and temperature are discussed in relation to predicted climate change.

Shade adaptation and shade tolerance in saplings of three Acer species from eastern North America.

Saplings of three, co-occurring maple species in a mature maple-beech forest differed in a suite of structural and physiological characters that separated the canopy species, Acer, saccharum, from the two subcanopy species, A. pensylvanicum and A. spicatum. Acer saccharum had both more dense wood and tougher and heavier but thinner leaves than the subcanopy species. Acer pensylvanicum had the largest, lightest leaves with high stomatal density and its canopy architecture was the most effective in terms of leaf display for light interception. Acer spicatum had weaker wood similar to that of A. pensylvanicum but also small, soft and relatively poorly displayed leaves. Both subcanopy species maintained marginally higher average rates of photosynthesis over the growing season in the understory environment. We consider juvenile A. saccharum only shade-tolerant, capable of persisting through long periods in the closed canopy until a gap occurs but not specifically adapted to the understory environment. Juvenile A. sacchrum appears to be constrained functionally by the requirements set by the canopy environment that adults will occupy. Characters such as high wood density are already expressed in the understory sapling; this investment in denser wood slows the growth of saplings, but is necessary for structural reasons in the adult. Juvenile A. saccaharum have morphological and photosynthetic characters better suited to gas exchange and extension growth under the increased photon flux densities in large forest gaps, characteristics that will also be advantageous in the sunlit canopy environment of adults.Both subcanopy maples appear to be more truly shade-adapted, although in somewhat different ways. Acer pensylvanicum has characteristics that enhance the potential for capture and utilization of sunflecks and is able to sustain higher growth rates than A. saccharum in the shaded subcanopy environment. Acer spicatum shares some shade-adapted features with A. pensylvanicum, and its habit of lateral spread through stem layering may confer an additional advantage in foraging for small light gaps.