What physiological processes permit insects to eat Eucalyptus leaves?

Many species of insects eat Eucalyptus foliage despite its relatively low nutritional value and the many plant secondary metabolites (PSMs) present, for example, terpenes, phenols and formylated phloroglucinols (FPGs). Formylated phloroglucinols are a new class of PSMs that act as antifeedants for possums and koalas. What physiological processes are present that permit insects to eat eucalypt foliage and how do PSMs influence insect feeding or digestion? Some trees seem to be repeatedly infested with eucalypt‐feeding insects, possibly as a result of previous chemosensory cues remaining from parental selection of a plant. Avoidance or storage of PSMs permit jarrah leafminers (Perthida glyphopa) and sawflies (Perga sp.) to consume eucalypt foliage without dealing with the majority of these compounds. Some PSMs can be metabolized by polysubstrate membrane oxidases as found in caterpillars or sawflies that feed on eucalypts. High midgut pH may be advantageous for nutrient extraction and PSM metabolism, and midgut pH ranges between 8.5 and 8.9 for caterpillars of Hyalarcta huebneri. Plant secondary metabolites may not be absorbed as a result of the combined presence of the peritrophic matrix and endogenous surfactants. Excretion of PSMs can be as metabolites or intact compounds. Both putative metabolites and sideroxylonal‐A, an FPG, are present in the faeces of larvae of the case moth, H. huebneri. The presence of sideroxylonal‐A in the food had an effect on the presence of 5‐hydroxytryptamine (5HT) in the central nervous system of caterpillars, as larvae fed leaves with a high concentration of sideroxylonal‐A had relatively more 5HT in the brain and central nervous system ganglia than larvae fed leaves containing a low concentration. Further work is necessary to clarify how PSMs are handled by eucalypt‐feeding insects and what effect FPGs have on feeding and digestion.     

Are compound leaves an adaptation to seasonal drought or to rapid growth? Evidence from the Amazon rain forest

Aim To assess the hypotheses that compound leaves of trees in the Amazon forest are an adaptation to drought and/or rapid growth. Location Amazon rain forest, South America. Methods Genera from 137 permanent forest plots spread across Amazonia were classified into those with compound leaves and those with simple leaves. Metrics of compound leaf prevalence were then calculated for each plot and regression models that accounted for spatial autocorrelation were used to identify associations between climate variables and compound leaf structure. We also tested for associations between compound leaf structure and a variety of ecological variables related to life history and growth strategies, including wood density, annual increase in diameter and maximum height. Results One plant family, Fabaceae, accounts for 53% of compound‐leaved individuals in the dataset, and has a geographical distribution strongly centred on north‐east Amazonia. On exclusion of Fabaceae from the analysis we found no significant support for the seasonal drought hypothesis. However, we found evidence supporting the rapid growth hypothesis, with possession of compound leaves being associated with faster diameter growth rates and lower wood densities. Main conclusion This study provides evidence that possession of compound leaves constitutes one of a suite of traits and life‐history strategies that promote rapid growth in rain forest trees. Our findings highlight the importance of carefully considering the geographical distribution of dominant taxa and spatial clustering of data points when inferring ecological causation from environment–trait associations.     

Use of leaves to inspect ectoparasites in wild chimpanzees: a third cultural variant?

We report 26 cases of using leaves as tools with which wild chimpanzees (Pan troglodytes schweinfurthii) in the Sonso community, Budongo Forest, Uganda, appeared to inspect objects removed during grooming. Careful removal of potential ectoparasites and delicate lip or manual placement on leaves followed by intense visual examination characterised this behaviour. It appears to be done to judge whether either ingestion or discarding is most appropriate, the former occurring in most cases. This behaviour may represent a third variant of ectoparasite handling, different from those described at Taï and Gombe, yet sharing features with the latter. These two East African techniques may thus have evolved from leaf grooming.     

Why some leaves are anthocyanic and why most anthocyanic leaves are red?

The adaptive significance of leaf reddening, as it occurs during specific developmental stages or after stress, has puzzled biologists for more than a century. Theoretically, the accumulation of a non-photosynthetic pigment competing with chlorophylls for photon capture would impose a photosynthetic cost, which should be paid off by the benefits afforded by anthocyanins under some circumstances. Hence, the proposed hypotheses presume protective functions against excess light, UV-B radiation, reactive oxygen species, water stress (osmoregulation) and herbivory. The existing arguments in favor of an anti-oxidant, anti-UV-B and osmoregulatory role are confounded by the co-occurrence in leaves of other compounds having the same properties, not absorbing visible light, attaining much higher concentrations and, in some cases, having a more appropriate location to fulfill the ascribed functions. Moreover, the excess light hypothesis should take into account that anthocyanins mainly absorb green photons, which are used photosynthetically in deeper cell layers needing less photoprotection. The more ecological, anti-herbivore hypotheses, consider red leaf color as a signal denoting high defensive commitment, as a camouflage obscuring the green reflectance indicative of a healthy leaf and/or as a device undermining the folivorous insects camouflage. The anti-herbivore hypotheses have not been thoroughly tested, yet they are compatible with the known optical preferences of insects and their underlying physiology. Overall, although a multiplicity of potential roles can be argued, the primary role may depend on the reference system, i.e. species, developmental stage or specific biotic and abiotic stressors.     

Desensitization to glucose 6-phosphate of phosphoenolpyruvate carboxylase from maize leaves by pyridoxal 5′-phosphate

Incubation of the nonphosphorylated form of maize-leaf phosphoenolpyruvate carboxylase (orthophosphate: oxaloacetate carboxy-lyase (phosphorylating), PEPC, EC 4.1.1.31) with the reagent pyridoxal 5′-phosphate (PLP) resulted in time-dependent, reversible inactivation and desensitization to the activator glucose 6-phosphate (Glc6P) and other related phosphorylated compounds. Both processes are not connected, since (i) when the PLP-modification was carried out in the presence of saturating ligands of the active site, which prevents inactivation, the desensitization to Glc6P is still observed, and (ii) under some experimental conditions the desensitization reaction is 4-times faster than the inactivation. Desensitization to Glc6P is first order with respect to PLP and has a second-order forward rate constant of 4.7±0.3 s⁻¹ M⁻¹ and a first-order reverse rate constant of 0.0046±0.0002 s⁻¹. Correlation studies between the remaining Glc6P sensitivity and mol of PLP residues incorporated per mol of enzyme subunit indicate that one lysyl group for enzyme monomer is involved in the sensitivity of the enzyme to Glc6P. The reactivity of this group is increased by polyethylene glycol and glycerol, while the reactivity of the lysyl group of the active site is not affected by these organic cosolutes. In the presence but not in the absence of the organic cosolutes, Glc6P by itself offers significant protection against desensitization, while increases the extent of inactivation. Free PEP or PEP-Mg have opposite effects, protecting the enzyme against inactivation and increasing the degree of desensitization. They also increases the protection against desensitization afforded by Glc6P. Finally, the PEPC inhibitor malate provides some protection against both inactivation and desensitization. Taken together, these results are consistent with PLP-modification of a highly reactive lysyl group at or near the allosteric Glc6P-site.     

Are red leaves photosynthetically active?

At irradiances close to those representing a sunny day, red and green leaves of poinsettia (Euphorbia pulcherrima) showed only minor differences in their photosynthetic capacities despite the strong differences in their pigment composition. However, contrarily to green leaves, red leaves did not show inhibition of photosynthesis at high irradiances, because anthocyanins protected chloroplasts from photoinhibition.     

Importance of ornithine-δ-aminotransferase to proline accumulation caused by water stress in detached rice leaves

Proline is synthesized either from glutamate or from ornithine in plants. Relatively little is known about the contribution of the pathway from ornithine to proline biosynthesis. In this paper we investigated the contribution of ornithine--aminotransterase (OAT), an enzyme responsible for ornithine pathway, to proline accumulation in water-stressed detached rice leaves. Although OAT activity increased with the increase of water stress duration, a pattern similar to that obtained for proline accumulation, the ornithine pathway in rice leaves seems to contribute little, if any, to proline accumulation under water stress condition. This conclusion was based on the observations that (a) gabaculine (50 M), an inhibitor of OAT, inhibited about 75% OAT activity caused by water stress but reduced only 20% of proline content and (b) cycloheximide, a protein synthesis inhibitor, had no effect on OAT activity induced by water stress but significantly reduced proline accumulation.     

Photosynthetic responses of sun- and shade-grown barley leaves to high light: is the lower PSII connectivity in shade leaves associated with protection against excess of light?

In this study, we have compared photosynthetic performance of barley leaves (Hordeum vulgare L.) grown under sun and shade light regimes during their entire growth period, under field conditions. Analyses were based on measurements of both slow and fast chlorophyll (Chl) a fluorescence kinetics, gas exchange, pigment composition; and of light incident on leaves during their growth. Both the shade and the sun barley leaves had similar Chl a/b and Chl/carotenoid ratios. The fluorescence induction analyses uncovered major functional differences between the sun and the shade leaves: lower connectivity among Photosystem II (PSII), decreased number of electron carriers, and limitations in electron transport between PSII and PSI in the shade leaves; but only low differences in the size of PSII antenna. We discuss the possible protective role of low connectivity between PSII units in shade leaves in keeping the excitation pressure at a lower, physiologically more acceptable level under high light conditions.     

Studies on Bisflavonóids of the leaves of Cephalotaxus fortunei Hook fvar alpina Native to China

Cephalotaxus fortunei Hook. f. var. alpina Li. is native to China. Two bisflavonoids were isolated and identified as sciadopitysis and amentoflavone-4',4'',7,7'-tetramethyl ether.     

Do high-tannin leaves require more roots?

The well-known deceleration of nitrogen (N) cycling in the soil resulting from addition of large amounts of foliar condensed tannins may require increased fine-root growth in order to meet plant demands for N. We examined correlations between fine-root production, plant genetics, and leaf secondary compounds in Populus angustifolia, P. fremontii, and their hybrids. We measured fine-root (<2mm) production and leaf chemistry along an experimental genetic gradient where leaf litter tannin concentrations are genetically based and exert strong control on net N mineralization in the soil. Fine-root production was highly correlated with leaf tannins and individual tree genetic composition based upon genetic marker estimates, suggesting potential genetic control of compensatory root growth in response to accumulation of foliar secondary compounds in soils. We suggest, based on previous studies in our system and the current study, that genes for tannin production could link foliar chemistry and root growth, which may provide a powerful setting for external feedbacks between above- and belowground processes.     

Species trait syndrome drives the leaves’ functional variations of dominant grasses to modifications in summer water supply

Climate change models predict a strong reduction of average precipitation, especially of the summer rainfall, and an increase in intensity and frequency of drought events in the Mediterranean region. The research aim was to understand how four dominant grass species (Arrhenatherum elatius, Cynosurus cristatus, Elymus repens, and Lolium perenne) in sub-Mediterranean meadows (central Apennines, Italy) modulate their resource acquisition and conservation strategies to short-term variation of the pattern of summer water supply. During summer 2016, using a randomized block design, we tested the effect of three patterns of summer water supply, differing in water amount and watering frequency, on leaf area, leaf dry mass, specific leaf area (SLA), leaf senescence, and plant height. Our results showed that dominant grass species can modulate their strategies to variation of the pattern of summer water supply, but the response of leaf traits and plant height is mediated by the set of functional characteristics of the species. E. repens and A. elatius, with summer green leaves, lower SLA, later flowering period, and deeper roots, were less influenced by changes in water amount. C. cristatus and L. perenne, which display acquisitive strategies (persistent leaves, higher SLA values), earlier flowering, and shallower roots were more influenced by changes in the pattern of summer water supply. Our results suggest that a short-term decrease in water availability might affect primarily species with trait syndromes less adapted to face summer drought.

     

Glycine decarboxylase is confined to the bundle-sheath cells of leaves of C3−C4 intermediate species

Immunogold labelling has been used to determine the cellular distribution of glycine decarboxylase in leaves of C₃, C₃–C₄ intermediate and C₄ species in the genera Moricandia, Panicum, Flaveria and Mollugo. In the C₃ species Moricandia foleyi and Panicum laxum, glycine decarboxylase was present in the mitochondria of both mesophyll and bundle-sheath cells. However, in all the C₃–C₄ intermediate (M. arvensis var. garamatum, M. nitens, M. sinaica, M. spinosa, M. suffruticosa, P. milioides, Flaveria floridana, F. linearis, Mollugo verticillata) and C₄ (P. prionitis, F. trinervia) species studied glycine decarboxylase was present in the mitochondria of only the bundle-sheath cells. The bundle-sheath cells of all the C₃–C₄ intermediate species have on their centripetal faces numerous mitochondria which are larger in profile area than those in mesophyll cells and are in close association with chloroplasts and peroxisomes. Confinement of glycine decarboxylase to the bundle-sheath cells is likely to improve the potential for recapture of photorespired CO₂ via the Calvin cycle and could account for the low rate of photorespiration in all C₃–C₄ intermediate species.Abbreviation and symbol kDa kilodaltons - CO₂ compensation point     

Why does Daphne pseudomezereum drop its leaves in the summer? An adaptive alternative to surviving forest shade

Daphne pseudomezereum A. Gray (Dpm) appears to be the only woody species in the north temperate forest that sheds its leaves in the summer while remaining green over winter (i.e. wintergreen leaf habit). Yet, the reason for this odd leaf habit has not been explored. To this end, we examined the microclimatic settings and ecophysiological traits of Dpm and its three native congeners in a field study of eight natural populations. In addition, we conducted a common garden experiment using Dpm plants where potential carbon gain across the seasons was estimated, using actual field microclimate data. Together, these data tested the hypothesis that Dpm retained traits of an open-grown upland ancestor, unable to adapt to the deep summer shade, it survived by becoming summer dormant and wintergreen. Our hypothesis was supported by patterns of leaf ecophysiological traits and carbon gain simulations in Dpm, consistent with the energetic feasibility of a summer dormancy followed by an autumn leaf sprout. We also conclude that carbon deficit driven by low light and high respiration cost is the trigger for the leaf habit of Dpm and assert that its phenological strategy represents a rare but viable alternative strategy for persistence in the temperate understory.     

Do small leaves expand faster than large leaves,and do shorter expansion times reduce herbivore damage?

Leaves are most vulnerable to herbivory during expansion. We hypothesised that one factor favouring small leaves could be that smaller-leaved species have shorter expansion times and are therefore exposed to high levels of herbivory for a shorter period than large leaves. In order to test this hypothesis, leaf expansion time and leaf area loss were measured for 51 species from Sydney, Australia. Strong positive correlations were found between leaf length and area and leaf expansion time, confirming that small leaves do expand in a shorter time than large leaves. The amount of leaf area lost was highly variable (from 0.5 to 90% of total leaf area), but was significantly related to both leaf expansion time and log leaf area. The amount of leaf area lost was not significantly correlated with specific leaf area nor with the presence of distasteful substances in the leaves, but was lower on species with hairy expanding leaves.     

Are fast growing birch leaves more asymmetrical?

Two recent papers (Martel et al. 1999, Lempa et al. 2000), linked rapid leaf growth in birches with high level of developmental instability, measured by leaf fluctuating asymmetry (FA). I discuss methodological approaches used in the referred papers and conclude that none of them provided incontrovertible support for the hypothesis on positive correlation between FA and leaf growth rate. A field study designed to test this hypothesis by an independent data set revealed no correlation between FA and either relative or absolute leaf growth rate in mountain birch either in within-tree or in between-tree comparisons. Character of the relationships between FA and growth may change with the index used to quantify developmental instability.