Effects of cotton condensed tannin on feeding and digestion in the cotton pest, Heliothis zea

Condensed tannin of cotton incorporated into an artificial diet caused growth inhibition in H. zea larvae whose susceptibility was dependent upon the concentration of the dietary tannin, and also upon the size of the feeding larvae.Larvae feeding on tannin-treated diet exhibited both decreased protease and invertase activities in the midgut caecal wall, as well as lowered total protein and sugar levels in the haemolymph as compared to controls. These differences, however, had no effect on assilimation and efficiency of conversion of digested matter into animal biomass and therefore may be secondary effects. Ultimately, inhibition of growth is attributed to a reduction in food consumption.     

Bitter problems in ecological feeding experiments: Commercial tannin preparations and common methods for tannin quantifications

In the field of plant–herbivore interactions, research methods where plant secondary metabolites are manipulated are becoming more and more popular. Among the most commonly used is tannic acid. However, recent studies have shown that different tannic acid preparations are not comparable in their tannin structures. While tannic acids are meant to contain only gallotannins (GTs), some commercial preparations compose mainly of more simple galloylglucoses (that have, e.g. much lower protein precipitation capacity than GTs) or even of gallic acid (the hydrolysis product of GTs). Another group of tannins used in feeding trials is condensed tannins (CTs), usually in the form of quebracho tannin. Quebracho, however, contains different CT structures than, e.g. leaves of many deciduous trees. Additionally, when analysed with the common acid-butanol assay for total CTs, quebracho tannins give even 30-fold lower absorbance than the CTs of those deciduous trees. In addition to above problems it has been shown that different tannins can give different response even within the same herbivore species, and that the same tannin structure can cause different response in different herbivores. Below we review these problems, as well as some means to deal with them.     

Hydrolyzable tannins as “quantitative defenses”: Limited impact against Lymantria dispar caterpillars on hybrid poplar

The high levels of tannins in many tree leaves are believed to cause decreased insect performance, but few controlled studies have been done. This study tested the hypothesis that higher foliar tannin levels produce higher concentrations of semiquinone radicals (from tannin oxidation) in caterpillar midguts, and that elevated levels of radicals are associated with increased oxidative stress in midgut tissues and decreased larval performance. The tannin-free leaves of hybrid poplar (Populus tremula × P. alba) were treated with hydrolyzable tannins, producing concentrations of 0%, 7.5% or 15% dry weight, and fed to Lymantria dispar caterpillars. As expected, larvae that ingested control leaves contained no measurable semiquinone radicals in the midgut, those that ingested 7.5% hydrolyzable tannin contained low levels of semiquinone radicals, and those that ingested 15% tannin contained greatly increased levels of semiquinone radicals. Ingested hydrolyzable tannins were also partially hydrolyzed in the midgut. However, increased levels of semiquinone radicals in the midgut were not associated with oxidative stress in midgut tissues. Instead, it appears that tannin consumption was associated with increased metabolic costs, as measured by the decreased efficiency of conversion of digested matter to body mass (ECD). Decreased ECD, in turn, decreased the overall efficiency of conversion of ingested matter to body mass (ECI). Contrary to our hypothesis, L. dispar larvae were able to maintain similar growth rates across all tannin treatment levels, in part, because of compensatory feeding. We conclude that hydrolyzable tannins act as “quantitative defenses” in the sense that high levels appear to be necessary to increase levels of semiquinone radicals in the midguts of caterpillars. However, these putative resistance factors are not sufficient to decrease the performance of tannin-tolerant caterpillars such as L. dispar.     

The role of phenolic compounds and other plant constituents in feeding by Canada geese in a coastal marsh

Summary We have assessed the relative importance of phenolic compounds, other secondary metabolites, and gross nutrient levels as feeding cues to Canada geese. Phenolic content was the most significant constituent influencing feeding selection by geese. Nutrient content had little or no effect on feeding selection.Correlative data showing the negative influence of plant phenolics on food choices by wild geese were supported by feeding preference tests. Extracts of unpalatable plants inhibited feeding by captive geese relative to extracts of palatable plants. In high phenolic plants, the phenolic containing methanol extract was more inhibitory than extractions made with petroleum ether. In a relatively low phenolic, unpalatable plant, an inhibitory factor was extractable in petroleum ether, indicating that for this species, another class of deterrents was involved. Preference tests with individual secondary metabolites showed that tannic acid and quebracho tannin were very effective in inhibiting feeding by geese and phenolic acids were slightly inhibitory, but a sesquiterpene lactone was not deterrent. These results point out the primacy of some secondary metabolites in determining food choices by geese.     

Effects of hydrolyzable and condensed tannin on growth and development of two species of polyphagous lepidoptera: Spodoptera eridania and Callosamia promethea

Summary The effects of tannins on survival, growth, and digestion were compared in two polyphagous species of Lepidoptera (one, the southern armyworm, a forb-feeder; and the other, the promethea silkmoth, a tree-feeder). Two different types of tannins (hydrolyzable and condensed) were incorporated into artificial basal diets in order to determine whether or not differential survival and growth would result between the forb feeder, which normally does not encounter tannins in its natural diet, and the tree-feeder, whose host species include many tanniniferous plants from several different families.Neonate larvae of the forb-feeding armyworms exhibited significantly suppressed 10-day growth rates at all tannin concentrations tested (0.5, 1.0, 1.5, 2.0, 2.5, and 3.0% of wet weight) for both the hydrolyzable and the condensed tannin compared to the control diet, however no dose-effect was detectable. In contrast, there were no detectable differences in neonate survival or growth through the first 10 days for the tree-feeding promethea silkmoth larvae fed diets with either tannic acid or quebracho tree condensed tannin.In order to determine the physiological mechanisms of action of these tannins against armyworms, we conducted detailed physiological bioassays of biomass and nitrogen utilization by penultimate instar larvae. Standard gravimetric feeding studies with both tannic acid and the quebracho tree condensed tannin demonstrated that reduced relative growth rates (RGR's) of Spodoptera eridania Cram. were due to the suppressed relative consumption rates (RCR's) and decreased conversion efficiencies (ECD's) rather than due to digestibility-reduction (as reflected by approximate digestibility, AD). As with the neonate larval growth rate suppression, there were no detectable dose responses at the different concentrations of tannic acid (0.25, 0.50, 0.75, 1.00, 2.50, and 5.0 percent) and condensed tannins from quebracho (0.25, 0.50, 0.75, 1.0, and 2.5 percent) in our penultimate instar studies.     

Relationships between leaf age and the food quality of cottonwood foliage for the gypsy moth,Lymantria dispar

Summary The cottonwood tree, Populus deltoides, continues to produce leaves late into the growing season, exposing midseason herbivores to leaves of a wide range of maturity. Gypsy moth larvae preferred and grew best on the oldest cottonwood leaves and suffered higher mortality and 85% less growth when fed young, expanding leaves. Concentration of phenolics in the youngest leaves was 3 times that in the oldest leaves and was negatively correlated with caterpillar growth rate. The active phenolics were not identified; tannin was present but its concentration changed more with season than leaf age.     

Effect of pine foliage damage on the incidence of larval diapause in the pine caterpillar Dendrolimus punctatus (Lepidoptera: Lasiocampidae)

The pine caterpillar Dendrolimus punctatus (Walker) with a larval facultative diapause is one of the most destructive insect pests of the pine tree Pinus massoniana in China. The larvae feeding on pine trees with different damage levels were studied to determine the induction of diapause under both laboratory and field conditions. Developmental duration of larvae before the third instar was the longest when fed with 75%–90% damaged needles, followed by 25%–40% damaged needles and intact pine needles, whereas mortalities did not differ among different treatments under the conditions of 25° and critical photoperiod 13.5:10.5 L:D. At 25°, no diapause was induced under 15:9 L:D, whereas 100% diapause occurred under 12:12 L:D regardless of the levels of needle damage. Incidences of larvae entering diapause when they were fed with intact, 25%–40% and 75%–90% damaged pine needles were 51.7%, 70.8% and 81% under 13.5:10.5 L:D, respectively. Similar results were obtained in the field experiment. Incidence of diapause was significantly different among the pine needle damage levels of pine trees when the photoperiod was close to the critical day length, indicating that the effect of host plants on diapause induction was dependent on the range of photoperiod. The content of amino acid and sugar decreased and tannin increased in pine needles after feeding by the pine caterpillars, suggesting that changed levels of nutrients in damaged needles or a particular substance emitted by damaged pine trees was perhaps involved in the diapause induction of the pine caterpillar.     

草地贪夜蛾在不同小麦品种上的取食选择性和适应性及其与叶片生化物质含量的关系

[目的]测定草地贪夜蛾Spodoptera frugiperda在不同小麦品种上的取食选择性及适应性,探索害虫为害与小麦品种间的关系,明确黄淮海麦区主栽小麦品种对草地贪夜蛾的抗性水平,为麦田抗虫品种布局和害虫综合防治提供理论依据.[方法]比较了草地贪夜蛾1和3龄幼虫对15个小麦品种的取食选择性,初步筛选出1和3龄幼虫取...     

The influence of addition of gallic acid, tannic acid, or quebracho tannins to alfalfa hay on in vitro rumen fermentation and microbial protein synthesis

Tannins may reduce rumen degradability of protein, increase the proportion of feed protein reaching the lower digestive tract for enzymatic digestion and thereby increase the efficiency of protein utilization. The objective was to assess the effects of different types and levels of tannins on rumen in vitro gas production and its kinetics, in vitro true degradability (IVTD) and rumen degradability of protein (IVRDP), and microbial protein synthesis by incubating alfalfa (Medicago sativa L.) hay in buffered rumen fluid. Alfalfa was incubated in buffered rumen fluid with and without the addition of different levels of gallic acid (GA), quebracho tannin (QT), or tannic acid (TA). Tannins at the lower inclusion levels had minimal effects on fermentation products compared to the higher levels. Addition of QT and TA reduced ammonium-N (NH₄⁺-N) concentration. Addition of QT at 20, 40, and 60 g/kg DM decreased NH₄⁺-N by 2, 7, and 12% compared with control whereas addition of TA reduced NH₄⁺-N by 5, 6, and 12% when added at 20, 40 and 60 g/kg DM, respectively. In experiment 2, addition of QT at 50, 100, and 150 g/kg DM, resulted in reduction of NH₄⁺-N by 12, 30, and 51%, respectively, compared with the control. Addition of TA at 50, 100, and 150 g/kd DM reduced NH₄⁺-N by 14, 26, and 47% compared with control. Inclusion of QT at 50, 100, 150 DM reduced IVRDP by 13, 30, and 36% compared with control whereas at these levels of inclusion, TA resulted in reduction of IVRDP by 14, 25, and 48%. Rate of gas production decreased (P<0.001), while asymptotic gas production increased (P<0.0001) with increasing level of GA and TA. Quebracho tannin decreased (P<0.0001) both the rate and asymptotic of gas production. Gallic acid had a positive effect on fermentation as indicated by increased gas production and total short chain fatty acids (SCFA) production. Quebracho tannin decreased 24 h gas production, IVTD, and total SCFA production. Acetate to propionate ratio increased with the addition of GA and but decreased when QT was added. Addition of tannins did not markedly increase total purines but numerical values tended to be higher in the presence of tannins compared with the control. Efficiency of microbial growth was lower in the presence of GA and unaltered by TA, but higher in the presence of QT compared with the control. The effect of tannins on rumen fermentation and protein degradation varied with type and level of tannins. In vivo studies will be conducted to validate the in vitro results.     

Condensed tannins in the diets of primates: a matter of methods?

To understand the ways in which condensed tannins (CT) affect primate diet selection and nutritional status, correct measurements are essential. In the majority of studies of the CT contents of primate foods, a tannin source such as "quebracho" is used to standardize CT assays, but the CT in quebracho tannin may not be similar to those in the plants of interest. We investigated how the choice of standard to calibrate CT assays affects the estimation of CT in the diets of mountain gorillas (Gorilla beringei). We purified the CT from gorilla foods and compared the actual amounts of CT in the foods with estimates produced by using the quebracho tannin. When quebracho was used, the estimates of CT contents of gorilla foods were, on average, 3.6 times the actual content of CT so that the amounts in frequently eaten gorilla foods were substantially overestimated. The overestimation for a given plant could not be predicted reliably and the ranking of plants by tannin content differed according to the standard used. Our results demonstrate that accurate measurements of CT necessitate the use of tannins purified from the plant species of interest. A reevaluation of primatology studies using interspecific comparisons of tannin content will provide new insights into primate food selection and nutritional ecology.     

Do birch-feeding caterpillars make the right feeding choices?

Summary The effects of leaf quality on caterpillar performance are frequently investigated, as are the feeding preferences of caterpillars, but rarely are the two considered together to see how preferences match performance. I looked at the preferences and performance of the specialist feeder Achlya flavicornis and the generalist Spilosoma luteum on Betula pendula leaves which were artificially damaged, mined or left undamaged, and on leaves of different ages. Often feeding preferences were not for the leaf types that gave faster development, larger pupae or greater survival. A. flavicornis preferred medium aged to older leaves though medium aged leaves produced smaller pupae. They did not discriminate between young and medium aged leaves, though larvae fed young leaves developed more slowly, and had lower survival and lower adult emergence. S. luteum larvae did not discriminate between damaged and undamaged leaves, though feeding on damaged leaves gave smaller pupae and lower adult emergence. I suggest that generalists like S. luteum might be unable to make feeding choices between leaves that are on the same tree and vary only slightly because populations are exposed to much greater variation across the wide range of potential food plants individuals may encounter. Possible differences in within-plant feeding selectivity between generalists and specialists should be investigated. A. flavicornis is a leaftier for which selection of a suitable leaf for tying could be more important in the first instance than feeding considerations. Larvae preferred younger to older leaves for tying, and larvae tended to feed on the tied leaves, so tying preferences explain the initial feeding preferences.     

Ecological tannin assays

Summary Estimations of condensed tannin content are generally based on calibration standard curves from Quebracho condensed tannins. We generated calibration standard curves from eight Sonoran Desert species for comparison with estimates of tannin concentrations derived from the Quebracho standard curve. Estimates of leaf tannin concentrations of each of the eight species using each species standard curve differed significantly with the estimates given by the Quebracho standard curve. Standard curves constructed from tannins from different individuals of three of the species varied significantly between, but not within, species. The efficiency of precipitation of protein bovine serum albumin (BSA) by each different tannin varied up to a factor of fifty for tannins of different species. Ordering species from highest to lowest based on tannin concentrations or binding efficiencies gave two different ranks. We argue that concentration or efficiency alone do not describe adequately tannin ecological activity. Instead, we suggest combining tannin concentrations and binding efficiencies to measure the protein precipitating potential of a leaf. Leaf protein precipitating potential is a more ecologically realistic parameter, we feel, for between-species comparisons than tannin content or binding efficiencies alone.Operated for the U.S. Department of Energy by the University of California under Contract No. DE-AC03-76-SF00012. This article was supported by the Director of the Office of Energy Research, Office of Health and Environmental Research     

The delta-endotoxin of Bacillus thuringiensis. V. On the occurrence of endotoxin fragments in hemolymph

Leucine-³H labeled crystals of Bacillus thuringiensis δ-endotoxin were fed to last-instar larvae of spruce budworm, Choristoneura fumiferana, eastern forest tent caterpillar, Malacosoma disstria, and silkworm, Bombyx mori. Radioactivity was detected in hemolymph 1 min after feeding in the first two species, but not until 3–5 min after feeding in silkworm larvae. Most of the radioactivity from hemolymph of all three species eluted from gel filtration columns at the same elution volume indicating similar moleculear weights (<1800 daltons).     

Influence of prey size on predation success by Zelus longipes L. (Het., Reduviidae)

The effects of prey size on the predatory responses of the reduviid Zelus longipes were studied through laboratory tests using larvae of the noctuid moth Spodoptera frugiperda as preys. In tests with one caterpillar, larvae of three different weight classes were offered individually to the predator. The prey weight was positively correlated with relative weight gain by the predator, mean feeding time and discarded biomass, but not with the relative extraction rate (defined as the relative weight gain by the predator by feeding time). The different sizes of caterpillars were attacked with the same frequency, but the successful attacks were more frequent in small larvae. The median mass of successfully attacked larvae was also less than that of unsuccessfully attacked. In tests with three caterpillars, larvae of three weight classes were offered at the same time; small caterpillars were more often attacked and killed than the medium and large ones. The results showed that even if larger preys resulted in more energy intake, when the choice is possible, smaller caterpillars were more likely to be attacked than medium and large. This is probably related to the fact that successful attacks were more frequent in small larvae, and also reduced the risk of injury to the predator.     

Captive roe deer (Capreolus capreolus) select for low amounts of tannic acid but not quebracho: fluctuation of preferences and potential benefits

Browsing ruminants have been shown to tolerate a certain amount of tannins in their natural diet, and preference trials with captive roe deer (Capreolus capreolus) have suggested an active selection for a low dose of hydrolysable tannins. In this study, we investigated the preference patterns for tannic acid, a source of hydrolysable tannins, and quebracho, a source of condensed tannins, in a series of preference trials with captive roe deer over time, using a pelleted feed that differed only in the respective tannin content. Additionally, two groups of four hand-raised roe deer fawns were fed either a control or a 3% tannic-acid containing diet and physiological parameters were compared after 7.5 months. There were large differences in preference patterns between the individual roe deer groups; quebracho was mostly avoided, whereas tannic acid was actively included in the diet in differing, low proportions. However, one group consistently preferred the quebracho diet over both the control or the tannic acid diet. For the tannic acid, the preference pattern often revealed an initial period of high preference, followed by a stable period of a moderate preference. The fawns on the tannic acid diet had a lower pellet intake and a higher relative mass gain than the fawns on the control diet; differences in salivary tannin-binding capacity and in blood antioxidant status were below significance. These results are the first indications of potential benefits of a low-dose tannin diet, which need further confirmation. The results of the preference trials demonstrate that the time pattern of tannin intake is not constant, and pose the question about the validity of short-term preference trials in general.     

Analysis of commercial proanthocyanidins. Part 1: the chemical composition of quebracho (Schinopsis lorentzii and Schinopsis balansae) heartwood extract

Quebracho (Schinopsis lorentzii and Schinopsis balansae) extract is an important source of natural polymers for leather tanning and adhesive manufacturing. We combined established phyto- and synthetic chemistry perspectives with electrospray mass spectrometry experiments to prove that quebracho proanthocyanidin polymers consist of an homologous series of flavan-3-ol based oligomers. The starter unit is always catechin which is angularly bonded to fisetinidol extender units. By comparison of the MS(2) fragmentation spectra of the oligomer with product ion scans of authentic catechin and robinetinidol samples, we proved that quebracho extract contains no robinetinidol, as is often reported. Quebracho proanthocyanidins have acid resistant interflavanyl bonds, due to the absence of 5-OH groups in fisetinidol, and the aDP cannot be determined via conventional thiolysis and phloroglucinolysis. We used the MS data to estimate a conservative (minimum value) aDP of 3.1.     

Tannin and tannate from the quebracho tree: an eco-friendly alternative for controlling marine biofouling

Current antifouling coatings are based on toxic compounds that can be harmful to the natural environment. A promising alternative to these compounds is the use of natural products that are non-toxic, but have antifouling properties. Tannins are natural, water-soluble, complex polyphenolic substances, which precipitate proteins and have anticorrosive and antimicrobial properties. In this study, the effect of quebracho tannin as a probable antifouling pigment in both laboratory and field trials is evaluated. As tannins have high solubility in aqueous media and consequently would leach rapidly, they were precipitated as aluminium tannate, which has an adequate solubility for use as a component in marine paints. In vitro exposure of Balanus amphitrite and Polydora ligni larvae to low concentrations of both quebracho tannin and saturated aluminium tannate solutions produced complete appendage immobilisation. In 28-d field trials of test gels, a significant decrease in micro- and macrofouling density and diversity in relation to the control gel was detected (p < 0.05). This study suggests that natural tannins could be employed as bioactive pigment for new antifouling technologies.     

The influence of phenolic compounds on the suitability of three Salix species as hosts for the willow beetle Phratora vulgatissima

The phenolic glycosides salicin and salicortin were found to influence larval growth and development rates and adult feeding preference of Phratora vulgatissima in laboratory feeding studies. Salicortin was more toxic to larvae than salicin, and none of the third instar larvae fed on Salix viminalis leaves amended with 1.52% (fresh mass) salicortin pupated. Condensed tannins (proanthocyanadins) did not affect larval performance. It was concluded that Salix burjatica resistance to willow beetle is due to the high levels of salicortin which occur in leaves of this species.     

Caterpillar assemblages on Chusquea bamboos in southern Ecuador: abundance,guild structure,and the influence of host plant quality

1. Information on the guild structure of foliage‐associated tropical insects is scarce, especially as caterpillars are mostly considered only as herbivores feeding on living leaves. However, many caterpillar species display alternative trophic associations, feeding on dead or withered leaves or epiphylls (‘non‐herbivores’). 2. To determine the contribution of these non‐herbivores, caterpillar communities associated with Chusquea Kunth (Poaceae) in the Andes of southern Ecuador were investigated. Caterpillars were collected at two elevation levels (montane rainforest ～2000 m and elfin forest at ～3000 m a.s.l.) and assigned to three feeding guilds (strict herbivores, non‐herbivores, and switchers) based on feeding trials. Foliage quality and leaf area were recorded to test for their influence on guild composition and caterpillar density. 3. Three hundred and eighty‐nine individuals belonging to 175 Lepidoptera species associated with Chusquea bamboos were found. The species richness of caterpillars was similarly high at both elevation levels but varied between feeding guilds. Approximately half (46.5%) displayed an alternative feeding association, i.e. were non‐herbivores (31.1%) or switchers (15.4%). 4. Caterpillar density was nearly two‐fold higher in the elfin forest, but only strict herbivores and switchers increased significantly with elevation. Leaf area positively influenced the density of strict herbivores and switchers; foliage quality only affected strict herbivores. The density of non‐herbivores did not differ significantly between the forest types and was not related to leaf area or foliage quality. 5. The present study underpins that non‐herbivores make up a considerable fraction of caterpillar communities in tropical mountain ecosystems and demonstrates that elevation, foliage quality and available plant biomass further shape feeding guild composition.     

Rainfall, nitrogen and host plant condition: consequences for the processionary caterpillar, Ochrogaster lunifer

1. At the end of November in subtropical areas of Australia, second-instar larvae of the processionary caterpillar Ochrogaster lunifer (Lepidoptera: Thaumetopoeidae) initiate feeding on the peripheral shoots of acacias (first-instar larvae do not feed). Field surveys at ten localities in south-east Queensland showed that larval survival was highly variable both among and within localities. Within-locality variation in larval growth was low compared with variation among localities. Larval growth and survival rates were higher at coastal and island localities, where November rainfall was high, than at drier inland localities.
2. Potted Acacia concurrens were grown in the greenhouse under high and low watering regimes, with and without nitrogen-rich fertilizer. Plant vigour (height, foliar water content and quantity of flush growth) was significantly greater in high-water treatments than in low-water treatments. Watering also affected foliar nitrogen, with plants in the high-water/no fertilizer treatment having similar nitrogen levels to those in fertilized treatments. Fertilizer increased foliar nitrogen levels of plants in low-water treatments and increased the number of shoots in high-water treatments. Different treatments had no effect on leaf toughness. After the first 3 weeks of feeding, size and survival of larvae were significantly reduced on the small, less vigorous plants in low water treatments. These results do not support the plant stress hypothesis.
3. Early-instar larvae (instar II–IV) developed more quickly and grew larger when reared on flush leaves than when reared on senescent leaves of A. concurrens . As water uptake affects the quantity of flush growth available to early stage larvae as well as foliar quality, rainfall and water availability may have important consequences for the distribution and population dynamics of the moth at local and regional scales.