African rainforest vegetation and rumen microbes: Phenolic compounds and nutrients as correlates of digestibility

Summary In order to refine hypotheses concerning food selection by generalist herbivores with ruminant-like digestive systems the chemical correlates of digestibility in such a system have been studied. Samples of seeds and leaves from tree species growing in two African rainforests (Douala-Edea Forest Reserve, Cameroon, and Kibale Forest, Uganda) were assayed for phenolic content and nutrient content, and in-vitro dry matter digestibility was analysed utilizing rumen inoculum from a fistulated steer. Both forests studied carry populations of colobine monkeys with ruminant like digestive tracts. Content of condensed tannins and, to a lesser extent of total phenolics, was found to be negatively correlated with digestibility; a result that may be attributable to the inctivation of microbial enzymes by tannins. The negative association of tannin content and digestibility was stronger in material from the Cameroon site, the vegetation of which contains considerably higher concentrations of tannins and is generally less digestible than that from the Ugandan site. Gross energy content of leaves was also found to be persistently negatively correlated with digestibility. The interpretation of this result is uncertain; however, gross energy yield may well reflect variation in content of cell wall polymers, especially lignin. For the complete set of data, tannins presented the strongest observed correlation with digestibility, but when only mature leaves were considered the relationship with gross energy appeared stronger. No strong association was noted between high nutrient content and high digestibility. This was attributed to the fact that the assay measured the extent of digestion under standardised and very favourable conditions of nutrient supply. The results obtained are discussed in relation to observations of leaf and seed selection preferences of Colobus spp. in these two forests.Publication 19-013 of the Wisconsin Regional Primate Research Center     

Foraging challenges of red colobus monkeys: influence of nutrients and secondary compounds

The diet selection of two groups of red colobus monkeys (Procolobus badius) in Kibale National Park, Uganda are considered with respect to protein, fiber, digestibility, alkaloids, total phenolics, tannins, saponins, and cyanogenic glycosides. Both groups selected young leaves over mature leaves and young leaves had more protein, were more digestible, and had a higher protein to fiber ratio than mature leaves. Young and mature leaves did not differ with respect to secondary compounds. There were no differences in the phytochemical factors examined between frequently eaten foods and leaves that red colobus were never known to eat, but were relatively common in the environment. Regression analyses predicting foraging effort from the phytochemical components of the large group's diet revealed selection for only one factor, foods that are high in protein and low in fiber, when differences in food tree availability were taken into consideration. A similar analysis with the small group did not suggest selection or avoidance of foods with respect to any of the factors considered. Previous studies have found the biomass of folivorous primates to be related to the ratio of protein to fiber concentration of mature leaves in the environment. These investigations have considered variation in folivore biomass and forest composition among sites separated by hundreds of kilometers; however, large variation in folivore abundance occurs over much smaller spatial scales. In Kibale National Park the average protein to fiber ratio of the mature leaves of the 20 most abundant tree species predicted the biomass of red colobus among four neighboring sites. We examined the generality of this relationship by adding our biomass and leaf chemistry values to previously published values; 62% of the variance in colobine biomass was explained by variation in the protein to fiber ratios of mature leaves at the sites. There was no evidence that red colobus avoided plants with high levels of secondary compounds. In fact, one of the most preferred trees (Prunus africana) was the species with the highest levels of cyanogenic glycosides, and the highest saponin levels were found in the young leaves of Albizia grandibracteata, the sixth and fourth most preferred plant species for the large and small groups, respectively.     

Food selection by black colobus monkeys (Colobus satanas) in relation to plant chemistry*

Black colobus monkeys (Golobus salanas) in the Douala-Edea Reserve, a rain-forest on the coast of Cameroon, have been shown to avoid young and mature leaves of most of the common plants in their habitat and to feed disproportionately heavily on leaves of rare plants. The proportion of leaves in the diet was low compared to most colobines studied, and the monkeys spent over half their feeding time eating seeds. Patterns of food selection were analysed in relation to distribution of nutrients, digestion–inhibitors and toxins in the vegetation. Colobw satanas select food items that are rich in mineral nutrients and nitrogen and low in content of the general digestion-inhibitors, lignin and tannin. They achieve this in the following ways: (i) by feeding preferentially on young leaves, which have higher nutrient content and lower contents of digestion-inhibitors than mature leaves; (ii) by eating those mature leaves with highest nument content relative to content of digestion-inhibitors; and (iii) by eating seeds, which are sources of readily available energy and which, as an item class, are less rich in digestion-inhibitors. Seeds at Douala-Edea appear to contain Ins nitrogen than leaves and C. satanas selects those seeds with highest nitrogen content. It is suggested that seed-feeding may be facilitated by the ability of the forestomach flora of these ruminant-like monkeys to detoxify some of the secondary compounds contained in seeds. Avoidance of most unused young and mature leaf items is correlated with a low nutrienudigestion-inhibitor ratio; avoidance of most unused seeds could be accounted for by their low nitrogen contents. Most items whose avoidance could not be explained in terms of these major constraints on food selection possess secondary compounds likely to be toxic. It is proposed that relative importance of digestion-inhibitors, low nutrient content and toxins as constraints on food selection by generalist herbivores will vary greatly among forests with different nutrient and secondary chemistry profiles.     

Tannin Dynamics of Propagules and Leaves of Kandelia candel and Bruguiera gymnorrhiza in the Jiulong River Estuary, Fujian, China

Changes in the total phenolics, condensed tannins (CT), protein-precipitable phenolics content and protein precipitation capacity were determined on a series of mangrove leaves from two true viviparous mangrove species (Kandelia candel and Bruguiera gymnorrhiza) at various stages of development and decomposition in the Jiulong River Estuary, Fujian, China. Similar measurements were also done for the propagules at different developmental stages. The results showed that the total phenolics, extractable condensed tannins, total condensed tannins, protein-precipitable phenolics content and protein precipitation capacity in young leaves were higher than those in mature and senescent leaves. Tannin dynamics during leaf decomposition varied with species, and the rapid loss of phenolics observed during decomposition can be ascribed to leaching and degradation. Protein-bound CT and fibre-bound CT tended to increase with leaf decomposition, with CT binding more strongly to protein than to fibre. Protein-bound CT was higher than fibre-bound CT with the exception of mature leaves. Total phenolics, extractable CT and protein-precipitable phenolics contents in flower tissues were relatively lower than those in hypocotyls at different developmental stages. Protein precipitation capacity fluctuated with the development of propagules. Increases in nitrogen in decaying litter, and declines in contents of total phenolics and total condensed tannins of detritus support the general conclusion that decomposing mangrove detritus can be a more palatable heterotrophic substrate than living leaves.     

A seasonal census of phenolics, fibre and alkaloids in foliage of forest trees in Costa Rica: some factors influencing their distribution and relation to host selection by Sphingidae and Saturniidae

The foliage of 80 species common in the Santa Rosa National Park, Costa Rica, has been analysed for content of total phenolics, condensed tannins, acid detergent fibre and water. Wherever possible analyses were performed at three stages in the life cycle of the leaf: young but fully expanded (coinciding with the beginning of the rainy season); middle-aged (two months later); and old (six months later). A comparison of the three age classes showed no significant change in the levels of phenolics or fibre as leaves aged but water content decreased significantly. A comparison of deciduous and evergreen species in the sample showed that the latter group had leaves with a significantly higher fibre content at all three sampling times, most particularly at the beginning of the rainy season, but other measures were not significantly different. Alkaloids were much more common in the foliage of deciduous species and it was observed that their distribution differed significantly from that of total phenolics and condensed tannins. It is suggested that the interaction that occurs between many tannins and alkaloids would be liable to reduce the defence capability of both classes of compounds if they occurred together.
High levels of defoliation occur in the early rainy season (third to tenth weeks) due to larvae of moths of the Sphingidae and Saturniidae. A comparison of investigated tree species that host larvae of these two taxa shows a striking dichotomy. Species that are selected by Sphingidae tend to be relatively deficient in levels of phenolics but are more likely to contain alkaloids, and probably other small toxic molecules. Saturniidae, on the other hand, appear to prefer host-species rich in phenolics but poor in alkaloids.     

Correlations between leaf toughness and phenolics among species in contrasting environments of Australia and New Caledonia

Background and Aims

Plants are likely to invest in multiple defences, given the variety of sources of biotic and abiotic damage to which they are exposed. However, little is known about syndromes of defence across plant species and how these differ in contrasting environments. Here an investigation is made into the association between carbon-based chemical and mechanical defences, predicting that species that invest heavily in mechanical defence of leaves will invest less in chemical defence.

Methods

A combination of published and unpublished data is used to test whether species with tougher leaves have lower concentrations of phenolics, using 125 species from four regions of Australia and the Pacific island of New Caledonia, in evergreen vegetation ranging from temperate shrubland and woodland to tropical shrubland and rainforest. Foliar toughness was measured as work-to-shear and specific work-to-shear (work-to-shear per unit leaf thickness). Phenolics were measured as ‘total phenolics’ and by protein precipitation (an estimate of tannin activity) per leaf dry mass.

Key Results

Contrary to prediction, phenolic concentrations were not negatively correlated with either measure of leaf toughness when examined across all species, within regions or within any plant community. Instead, measures of toughness (particularly work-to-shear) and phenolics were often positively correlated in shrubland and rainforest (but not dry forest) in New Caledonia, with a similar trend suggested for shrubland in south-western Australia. The common feature of these sites was low concentrations of soil nutrients, with evidence of P limitation.

Conclusions

Positive correlations between toughness and phenolics in vegetation on infertile soils suggest that additive investment in carbon-based mechanical and chemical defences is advantageous and cost-effective in these nutrient-deficient environments where carbohydrate may be in surplus.Key words: Antiherbivore defence, leaf toughness, mechanical defence, chemical defence, phenolics, trade-offs     

Diets of two sympatric colobines in Zaire: More evidence on seed-eating in forests on poor soils

Recent studies have shown that the so- called folivorous African colobines eat a significant amount of seeds. There is disagreement as to whether seed-eating is due to the poor quality of tree foliage, due in turn to poor soils, or to the fact that seeds are a normal part of colobine diets. To test these hypotheses, we studied feeding of red colobus monkeys, Colobus badius tholloni,and black-and-white colobus monkeys, Colobus angolensis angolensis,in a tropical rain forest of Central Zaire (Salonga National Park). We conducted studies on soil properties, vegetation composition, and the availability and chemical contents of food plants simultaneously. Soils were very acid, with a high percentage of sand, very low cation exchange capacity, and very low exchangeable bases. The forest was dominated by legumes (45.6% of trees), among which the Caesalpinioideae were the best represented (85%). C. badiusfed mostly on leaves (61%) and seeds (33%), legumes making up 65% of their diet. C. angolensisfed mostly on seeds (50%) and leaves (27%); 39% of their diet came from legumes. The two species tended to select items richer in crude protein or lipids or both. Total phenolics and condensed tannins were abundant in the foliage and seeds but were poor predictors of colobine choice of food. Intersite comparisons show that colobines in Zaire ate a higher proportion of seeds than all other related species so far studied in Africa and that the Salonga forest had among the poorest soils and harbored the highest percentage of Caesalpinioideae. Our results confirm that seed- eating is more common among colobines living in areas where soils are poor. They strongly suggest that this link is mediated through forest composition, especially the abundance of legumes, and that the development of seed- eating results both from the high availability of nutrient-rich seeds and from the poor quality of mature tree foliage.     

Variation in defence strategies in two species of the genus Beilschmiedia under differing soil nutrient and rainfall conditions

The relationships between various leaf functional traits that are important in plant growth (e.g., specific leaf area) have been investigated in recent studies; however, research in this context on plants that are highly protected by chemical defences, particularly resource-demanding nitrogen-based defence, is lacking. We collected leaves from cyanogenic (N-defended) Beilschmiedia collina B. Hyland and acyanogenic (C-defended) Beilschmiedia tooram (F. M. Bailey) B. Hyland at high- and low-soil nutrient sites in two consecutive years that varied significantly in rainfall. We then measured the relationships between chemical defence and morphological and functional leaf traits under the different environmental conditions. We found that the two species differed significantly in their resource allocation to defence as well as leaf morphology and function. The N defended species had a higher leaf nitrogen concentration, whereas the C-defended species had higher amounts of C-based chemical defences (i.e., total phenolics and condensed tannins). The C-defended species also tended to have higher force to fracture and increased leaf toughness. In B. collina, cyanogenic glycoside concentration was higher with higher rainfall, but not with higher soil nutrients. Total phenolic concentration was higher at the high soil nutrient site in B. tooram, but lower in B. collina; however, with higher rainfall an increase was found in B. tooram, while phenolics decreased in B. collina. Condensed tannin concentration decreased in both species with rainfall and nutrient availability. We conclude that chemical defence is correlated with leaf functional traits and that variation in environmental resources affects this correlation.     

Post-cultivation Secondary Succession in a Venezuelan Lower Montane Rain Forest

Since tropical rain forests are widely threatened by conversion to agriculture, even within protected areas, an understanding of recovery processes is important for restoration of forest ecosystems and thus conservation of their biodiversity. Secondary succession following land clearance and crop cultivation was studied in a lower montane rain forest in a protected area of the Venezuelan Cordillera de la Costa Central. Forest recovery was studied using a chronosequence of eight 20 × 20 m plots which represented four forest types ca.10 year-old Secondary Forest, ca. 20 year-old Secondary Forest, ca. 35 year-old (uncultivated) secondary forest and mature forest. Species richness and structural complexity increased during succession, with the oldest secondary forest having a physiognomy comparable to the mature forest. Species diversity was lower in the secondary forests than the mature forest, and their floristic composition was distinct. Four phases are hypothesized to occur in the succession process, each with a distinctive species assemblage: initial colonisation by non-woody vegetation; establishment and canopy closure by short-lived small-seeded woody pioneer species; replacement by longer-lived secondary species; and gradual replacement by mature forest large-seeded climax species. Full recovery of the forests in the protected area is likely to take many years, although it may be assisted through conservation management measures.     

The Herbivore Assemblage, Herbivory and Leaf Chemistry of the Mangrove Kandelia obovata in Two Contrasting Forests in Hong Kong

The herbivore assemblage, intensity of herbivory and factors determining herbivory levels on the mangrove Kandelia obovata (previously K. candel, Rhizophoraceae) were studied over a 13-month period at two forests with contrasting growing conditions in Hong Kong. Mai Po was part of an eutrophic embayment in the Pearl River estuary and generally offered more favourable conditions for mangrove growth, whereas Ting Kok had a rocky substratum and oceanic salinity. Twenty-four insect herbivore species were recorded on K. obovata, with lepidopteran larvae that consume leaf lamina being the dominant species. While leaf litter production was similar at the two forests, herbivory level at Mai Po (mean = 3.9% in terms of leaf area loss) was more severe than that at Ting Kok (mean = 2.3%). Peak herbivory levels were found in summer at both locations (6.5% for Mai Po and 3.8% for Ting Kok). Young leaves of K. obovata at both locations were generally preferred by the herbivores from the period of late spring to summer. Concentrations of most feeding deterrents (ash, crude fibre, and total soluble tannins) were significantly higher in both young and mature leaves at Ting Kok, whereas leaf nutrients (total nitrogen and water) were the same at the two sites. Young leaves at Ting Kok contained about 30% more tannins than their counterparts at Mai Po. Significant differences in leaf chemistry also existed between young and mature leaves at either site. The differences were concomitant with the observed patterns of leaf herbivory on K. obovata, and suggest a potential relationship between environmental quality and plant defence against herbivory.     

Defensive traits of savanna trees – the role of shoot exposure to browsers

To investigate patterns of plant defence, a study was conducted on 13 tree species in Botswana. We tested the hypotheses that (1) shoots on small, young trees are more defended than shoots at the same height on tall, mature individuals and (2) shoots within browsing height are more defended than shoots above the reach of browsers, on mature trees. Plant traits studied were total phenolics, tannin activity, nitrogen (N), acid-detergent fibre (ADF), and degree of spinescence (in three species). We found some support for the first hypothesis. Overall young trees had a significantly higher concentration of total phenolics than mature trees and on species level these differences were significant for three of the 13 species. However, we found no differences in tannin activity, spinescence or ADF concentration between young and mature trees, and four species had higher levels of N in young trees, suggesting higher overall palatability. Contrary to our second hypothesis, shoots within browsing height were less chemically defended (total phenolics and tannin activity) than shoots above browsing height in six of 13 species. Two species had less ADF in lower shoots, and no significant differences were found in N or spinescence for any species. This study included a broad range of tree species at two sites and the results indicate that general theories of plant defence, originating from boreal and northern temperate forests, cannot be simply applied to savanna ecosystems.     

Rethinking the role of many plant phenolics – protection from photodamage not herbivores?

Phenolics have been considered classic defence compounds for protecting plants from herbivores, ever since plant secondary metabolites were suggested to have evolved for that reason. The resource availability and carbon-nutrient balance hypotheses proposed that variation in phenolic levels between and within plant species reflects environmental availability of nutrients and light, and represents a trade-off in allocation by plants to growth and defence against herbivores. In contrast to these concepts, we suggest that (1) the main role of many plant phenolics may be to protect leaves from photodamage, not herbivores; (2) they can achieve this by acting as antioxidants; and (3) their levels may vary with environmental conditions in order to counteract this potential photodamage. We therefore suggest that patterns in phenolic levels, often used to support the concept of trade-off between growth and herbivore defence in relation to resource availability, may actually reflect different risks of photodamage. We suggest that the level of many phenolics is low under some environmental conditions, not because resources to produce them are limited, but simply because the risk of photodamage is low and they are not required. If our photodamage hypothesis is correct, a reassessment of the ecological and evolutionary role of many phenolics in plant defence theory is required.     

The diet of chimpanzees in the Budongo Forest Reserve, Uganda

Chimpanzee diets are highly variable, but in all cases dominated by ripe fruit. Previous work has suggested that in the Budongo Forest Reserve, Uganda, ripe fruit remains available throughout the year, but dietary data for chimpanzees at this site remain sparse. This paper details the diet of a recently habituated community of Budongo chimpanzees living in the Sonso region. Chimpanzees of this community spent 64.5% of their feeding time eating fruit, and 19.7% eating arboreal leaves. Terrestrial herbaceous vegetation formed only a minor component of the diet. In these general dietary characteristics, Budongo chimpanzees show greater similarity to those of the Gombe National Park than they do to those in the geographically closer and floristically more similar Kibale Forest National Park. The diet was dominated by the fruit of four species and leaves of two species, although the composition of the diet varied from month to month, remaining diverse. Figs were consumed throughout much of the year, and in consequence, should perhaps be regarded as a staple, rather than fallback, food. A period of food scarcity was not apparent in this study and future comparative nutritional investigation will be required to determine whether these chimpanzees face a time of dietary hardship.     

Effects of artificial grassland establishment on soil nutrients and carbon properties in a black-soil-type degraded grassland

Despite a growing knowledge of nutrient limitation for mangrove species and how mangroves adapt to low nutrients, there is scant information about the relative importance of N:P ratio and leaf phenolics variability in determining nutrient conservation. In this study, we evaluated possible nutrient conservation strategies of a mangrove Rhizophora stylosa under nutrient limitation. 1. The leaf nutrient concentrations of R. stylosa changed with season, with the highest N concentration in winter and the highest P concentration in spring for both mature and senescent leaves. Leaf N and P concentrations decreased significantly during leaf senescence. Based on N:P ratios R. stylosa forest was N-limited. Accordingly, the nitrogen resorption efficiency (NRE) was significantly higher than phosphorus resorption efficiency (PRE) for the R. stylosa leaves during leaf senescence. The NRE and PRE both reached the highest in the autumn. Average N and P concentrations in the senescent leaves were 0.15% and 0.06% for R. stylosa, respectively, indicating a complete resorption of N and an incomplete resorption of P. There was a significant negative correlation between nitrogen resorption proficiency (NRP) and NRE, meanwhile phosphorus resorption proficiency (PRP) and PRE correlation was also highly significantly. 2. R. stylosa leaves contained relatively high tannin level. Total phenolics, extractable condensed tannins and total condensed tannins contents increased during leaf senescence, and changed between seasons. The lowest concentrations of total phenolics, extractable condensed tannins and total condensed tannins occurred in summer, total phenolics concentrations were inversely related to nitrogen or phosphorus concentrations. 3. Our results confirmed that resorption efficiency during leaf senescence depends on the type of nutrient limitation, and NRE was much higher than PRE under N-limited conditions. R. stylosa forest developed several nutrient conservation strategies in the intertidal coastline surroundings, including high nitrogen resorption efficiency, low nutrient losses and high tannins level.     

Tannins and nitrogen dynamics in mangrove leaves at different age and decay stages (Jiulong River Estuary,China)

Changes in the total phenolics, condensed tannins (CT), protein-precipitable phenolics content and protein precipitation capacity were determined on mangrove leaves representing a range of maturation and senescence in the Jiulong River Estuary, Fujian, China. The results showed that the total phenolics, extractable condensed tannins, total condensed tannins, protein-precipitable phenolics content and protein precipitation capacity in young leaves were higher than those in mature and senescent leaves. With leaf maturation and senescence, total pehnolics decreased. The rapid loss of phenolics observed following senescence and abscission can be ascribed to leaching and degradation. Protein-bound CT and fibre-bound CT tended to increase with leaf maturity, and decreased with senescence. Protein-bound CT and fibre-bound CT increased with leaf decomposition, with CT binding more strongly to protein than to fibre. The increases in nitrogen contents and ash free caloric values, and declines in total phenolics and total condensed tannins of mangrove leaf litter suggests that partially decomposed mangrove detritus is a palatable heterotrophic substrate, and thus may be an important source of matter and energy for the estuarine food web. Handling editor: K. Martens     

东莞大屏嶂森林公园的现状植被及林分改造对策

采用改进的线路调查方法对东莞大屏障森林公园植被进行调查。结果表明大屏障森林公园植物种类多样,植物资源丰富。公园的现状植被可以划分为5个类型,8个亚型。森林可分为次生常绿阔叶林、人工马占相思林Acacia mangium、油茶林Camellia oleifera等3个亚型。此外,还有荒草坡、农田植被以及其他类型的植被。据此探讨了对大屏障森林公园植被的改造对策,为森林公园的建设提供依据。     

Tannin allocation in blackbrush (Coleogyne ramosissima)

Blackbrush (Coleogyne ramosissima) is a spinescent shrub occurring in nearly monospecific stands in the southwestern United States. Winter browsing by domestic goats stimulates spring twig production by blackbrush plants. Current season's twigs produced by older branches growing on the outer edges of the plant canopy (terminal branches) contained 2.37 times more tannins than did older twigs. Within blackbrush plants, current season's twigs on terminal branches contained 1.34 times more tannins than did current season's twigs on sprouts and younger branches (basal branches). When analysed separately, current season's terminal twigs contained 1.47 times more tannins than did leaves, while current season's basal twigs contained 1.10 times more tannins than did leaves. Phytochemicals, mainly phenolics, in blackbrush reduce its risk of being over-utilized by domestic herbivores, and the occurrence and distribution of tannins in blackbrush lend support to hypotheses dealing with secondary compounds as plant defences.     

Phenolic and mineral content of leaves influences decomposition in European forest ecosystems

Summary Factors influencing decomposition in European forests growing on different soils were studied in stands dominated by the European beechFagus sylvatica L. Phenolic contents of freshly fallen leaves ofF. sylvatica growing on nutrient-poor soils (acid sandy soil) were higher than those of similar leaves on nutrient-rich soils (calcareous mull soil). Analysis of fallen leaves of different ages showed rapid decay of phenolics during the first winter on the ground. After 1 year the phenolic content of leaves ofF. sylvatica growing on nutrient-poor soils was still twice as high as in similar leaves on nutrient-rich soils. Field and laboratory experiments showed that a major decomposer (Oniscus asellus, Isopoda) preferred leaves from trees on nutrient-rich soils. Mineral contents of leaves ofF. sylvatica growing on different soils differed: on rich soils leaves had higher contents of Ca, Mg, Na, and K. These elements are important nutrients for decomposers. The distribution of major decomposers reflects the mineral content of their diet, which in turn reflects soil type. Different rates of leaf turnover and nutrient turnover in different forest ecosystems (even when the same tree species is dominant) are due to the decomposing system, which is influenced by the phenolic and mineral contents of the leaves.     

Growth responses of gypsy moth larvae to elevated CO2: the influence of methods of insect rearing

Abstract The effects of elevated CO₂ on foliar chemistry of two tree species (Populus pseudo‐simonii Kitag. and Betula platyphylla) and on growth of gypsy moth (Lymantria dispar L.) larvae were examined. Furthermore, we focused on the comparison of results on the growth responses of larvae obtained from two methods of insect rearing, the no‐choice feeding trial performed in the laboratory or in situ in open‐top chambers. On the whole, both primary and secondary metabolites in the leaves of the two tree species were significantly affected by main effects of time (sampling date), CO₂ and species. Elevated CO₂ significantly increased the C : N ratio and concentrations of the soluble sugar, starch, total nonstructural carbohydrates, total phenolics and condensed tannins, but significantly decreased the concentration of nitrogen. Higher contents of total phenolics and condensed tannins were detected in the frass of larvae reared in elevated CO₂ treatments. Overall, the growth of gypsy moth larvae were significantly inhibited by elevated CO₂ and CO₂‐induced changes in leaf quality. Our study did not indicate the two methods of insect rearing could influence the direction of effects of elevated CO₂ on the growth of individual insects; however, the magnitude of negative effects of elevated CO₂ on larval growth did differ between the two insect rearing methods, and it seems that the response magnitude was also mediated by larval age and host plant species.     

Quantitative changes in secondary metabolites of dark-leaved willow (Salix myrsinifolia) exposed to enhanced ultraviolet-B radiation

This is a study of the impact of increased ultraviolet-B (UV-B) radiation on the secondary chemistry of Salix myrsinifolia (dark-leaved willow). For nearly two decades, the loss of stratospheric ozone above the high latitudes of the Northern Hemisphere has increased UV-B radiation (280–320 nm) over the long-term mean. Willows (Salicaceae) are widely distributed in these northern regions. To determine the effects of increased UV-B radiation on willows, the plantlets of three clones of S. myrsinifolia were grown under ambient (3.6 kJ m⁻² day⁻¹) or enhanced (7.18 kJ m⁻² day⁻¹) UV-B irradiance. After the 2-week indoor experiment, the concentrations of UV-B-screening phenolics (flavonoids and phenolic acids) and low-UV-B-screening phenolics (salicylates and condensed tannins) in fresh leaves were investigated and the biomass of leaves, stems and roots was determined. As expected, the total amount of flavonoids in willow leaves clearly increased when plantlets were exposed to higher UV-B irradiation. However, the degree of increase of individual compounds varied: luteolin-7-glucoside, monomethyl-monocoumaryl-luteolin-7-glucoside and one myricetin derivative increased significantly, while the apigenin-7-glucuronide increased only slightly. The enhanced UV-B also increased the amount of p -hydroxycinnamic acid derivative. The UV-B effects on other phenolic acids and tannins were minor. In contrast to the other phenolics, the amounts of two salicylates, salicin and saligenin, decreased under enhanced UV-B irradiation. Our results indicate that the concentrations of both UV-B-screening and low-UV-B-screening phenolic compounds in leaves of S. myrsinifolia may vary in response to elevated UV-B radiation. However, while the UV-B protective flavonoids and phenolic acids accumulate during UV-B exposure, the concentrations of certain salicylates decrease.