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William J Foley

Publications and source records attributed to William J Foley.

12 recordsLinked to original sources

Dugong grazing and turtle cropping: grazing optimization in tropical seagrass systems?

Grazing by dugongs and cropping by green turtles have the capacity to alter the subsequent nutritional quality of seagrass regrowth. We examined the effects of simulated light and intensive grazing by dugongs and cropping by turtles on eight nutritionally relevant measures of seagrass chemical composition over two regrowth periods (short-term, 1-4 months; long-term, 11-13 months) at two seagrass communities (a mixed species community with Zostera capricorni, Halophila ovalis, Halodule uninervis, Cymodocea rotundata and C. serrulate; and a monospecific bed of Halodule uninervis) in tropical Queensland, Australia. The concentrations of organic matter, total nitrogen, total water-soluble carbohydrates, total starch, neutral detergent fiber, acid detergent fiber, acid lignin, as well as the in vitro dry matter digestibility (IVDMD) were measured in the leaves and below-ground parts of each species using near-infrared reflectance spectroscopy (NIRS). Regrowth of preferred species such as H. ovalis and H. uninervis from simulated intensive dugong grazing after a year exhibited increased (by 35 and 25%, respectively, relative to controls) whole-plant N concentrations. Similarly, regrowth of H. ovalis from simulated turtle cropping showed an increase in the leaf N concentration of 30% after a year. However, these gains are tempered by reductions in starch concentrations and increases in fiber. In the short-term, the N concentrations increased while the fiber concentrations decreased. These data provide experimental support for a grazing optimization view of herbivory in the tropical seagrass system, but with feedback in a different manner. Furthermore, we suggest that in areas where grazing is the only major source of natural disturbance, it is likely that there are potential ecosystem level effects if and when numbers of dugongs and turtles are reduced.

Animals↗

The detoxification limitation hypothesis: where did it come from and where is it going?

The detoxification limitation hypothesis is firmly entrenched in the literature to explain various aspects of the interaction between herbivores and plant toxins. These include explanations for the existence of specialist and generalist herbivores and for the prevalence of each of these. The hypothesis suggests that the ability of mammalian herbivores to eliminate plant secondary metabolites (PSMs) largely determines which plants, and how much, they can eat. The value of the hypothesis is that it provides a clear framework for understanding how plant toxins might limit diet breadth. Thus, it is surprising, given its popularity, that there are few studies that provide experimental support either for or against the detoxification limitation hypothesis. There are two likely reasons for this. First, Freeland and Janzen did not formally propose the hypothesis, although it is implicit in their paper. Second, it is a difficult hypothesis to test, requiring an understanding of the metabolic pathways that lead to toxin elimination. Recent attempts to test the hypothesis appear promising. Results suggest that herbivores can recognize mounting saturation of a detoxification pathway and adjust their feeding accordingly to avoid intoxication. One strategy they use is to ingest a food containing a toxin that is metabolized by a different pathway. This demonstrates that careful selection of food plants is a key to existing in a chemically complex environment. As more studies characterize the detoxification products of PSMs, we will better understand how widespread this phenomenon is.

Food Preferences↗

Conflicting demands on detoxification pathways influence how common brushtail possums choose their diets.

Most herbivores eat more and survive better when they have access to a variety of foods. One explanation involves the detoxification of plant secondary metabolites (PSMs). By feeding from a variety of plants that contain different classes of PSMs, animals can use multiple detoxification pathways and presumably consume more food. Although popular, this theory is difficult to test because it requires knowledge of the detoxification pathways of each PSM in the diet. We established that common brushtail possums (Trichosurus vulpecula) use various combinations of oxidation, hydrolysis, and conjugation with glucuronic acid (GA) or glycine to detoxify six PSMs. Compared to their ingestion of a single PSM, possums ate more when offered a choice between two diets containing PSMs that require apparently independent detoxification pathways (benzoate and 1,8-cineole, benzoate and p-cymene, benzoate and orcinol, benzoate and salicin, or orcinol and 1,8-cineole). However, possums still did not eat as much of these diets as they did of a basal diet free of PSMs. This suggests that detoxification pathways are never independent, but are separated instead by degrees. In contrast, possums offered a choice of two PSMs that require competing detoxification pathways (1,8-cineole and p-cymene, 1,8-cineole and salicin, or orcinol and salicin) ate no more than when offered diets containing one of the compounds. There was an exception: even though both rutin and orcinol are detoxified via conjugation with GA, the feeding behavior of possums did not suggest competition for detoxification pathways. This implies that the supply of GA is not limiting. This study provides the first convincing evidence that herbivorous mammals can eat more by selecting mixed diets with a diversity of PSMs that make full use of their detoxification potential. It also emphasizes that other behavioral and physiological factors, such as transient food aversions, influence feeding behavior.

Animals↗

Sideroxylonal in Eucalyptus foliage influences foraging behaviour of an arboreal folivore.

Plant secondary metabolites (PSMs) offer plants chemical defences against herbivores, and are known to influence intake and diet choice in both insect and mammalian herbivores. However, there is limited knowledge regarding how PSMs influence herbivore foraging decisions. Herbivore foraging decisions, in turn, directly impact on which individual plants, and plant species, are selected for consumption. We took advantage of the natural variation in sideroxylonal concentrations in the foliage of Eucalyptus melliodora (Cunn. ex Schauer) to investigate feeding patterns of a marsupial folivore, the common ringtail possum, Pseudocheirus peregrinus (Boddaert 1785). Foliage, collected from six trees, contained between 0.32 and 12.97 mg g-DM(-1) sideroxylonal. With increasing sideroxylonal concentrations, possums decreased their total intake, rate of intake and intake per feeding bout, and increased their cumulative time spent feeding. Possums did not alter their total feeding time, number of feeding bouts or time per feeding bout in response to increasing sideroxylonal concentrations. Results demonstrate important behavioural changes in foraging patterns in response to sideroxylonal. These behavioural changes have important implications, in relation to altered foraging efficiency and potential predation risk, for herbivores foraging in the field. As a result, the spatial distribution of dietary PSMs across a landscape may directly influence herbivore fitness, and ultimately habitat selection of mammalian herbivores.

Animals↗

Tree use by koalas in a chemically complex landscape.

Although defence against herbivores is often argued to be the main action of plant secondary metabolites (PSMs), very few examples have demonstrated that intraspecific variation in PSM concentrations influences foraging by wild vertebrate herbivores. Experiments with captive animals often indicate that PSM concentrations influence how much herbivores eat from individual plants, but these experiments do not replicate the subtle trade-offs in diet selection faced by wild animals, which must avoid predators and extremes of weather, interact with conspecifics, and achieve a balanced, nutritious diet, while avoiding intoxication by PSMs. We characterized the foliar chemistry of every tree from two Eucalyptus species available to a population of koalas (Phascolarctos cinereus) and considered rates of tree visitation over a ten-year period. We show that visitation rate was most strongly influenced by tree size, but that koalas also visited trees less frequently if the foliage contained either high concentrations of deterrent PSMs known as formylated phloroglucinol compounds, or low concentrations of nitrogen. Consequently, plant chemistry restricts the use of trees by this herbivore, and thus limits the food available to koalas and potentially influences koala populations.

Animals↗

Eucalyptus foliar chemistry explains selective feeding by koalas.

The koala is the quintessential specialist herbivore, feeding almost exclusively on Eucalyptus foliage. Consequently, the limitations imposed on the koala's diet by plant defences indicate the extent to which evolutionary adaptations allow mammalian herbivores to circumvent such defences. We tested whether a recently discovered group of plant secondary metabolites, the formylated phloroglucinol compounds (FPCs), deters koalas from feeding on some eucalypt foliage. We found that captive koalas ate less foliage in a single night from trees with high FPC concentrations. Individual trees also differ in the types of FPC they possess, but for a given eucalypt species, most FPCs were similarly effective deterrents. Two closely related and sympatric eucalypt species could be clearly separated by the amounts that koalas ate from each; however, this difference could not be explained by total FPC concentrations alone. We suggest, that in this case, the presence of a distinct type of FPC deters koala herbivory on the less palatable species, and may have facilitated the evolutionary divergence of these species. We conclude that plant defences probably play an important role in determining the distribution and abundance of koalas.

Animals↗

Plant secondary metabolites and vertebrate herbivores--from physiological regulation to ecosystem function.

Plant secondary metabolites can constrain the diet of vertebrates and these effects can flow through to community dynamics. Recent studies have moved beyond attempting to correlate diet choice with secondary metabolite profiles and instead focus on mechanisms that animals use to detect toxins and to regulate their intake and absorption. These include molecularly determined taste specificity, serotonin-mediated learning and the control of toxin absorption by permeability-glycoproteins. Focus on the detoxification pathways employed by specialist and generalist herbivores has facilitated explicit tests of the long-standing hypothesis that detoxification rates limit feeding. Understanding the molecular basis of differences amongst species in their tolerance of plant secondary metabolites opens many opportunities for understanding the evolutionary history of interactions between vertebrates and their food plants.

Animals↗

Jensenone: biological reactivity of a marsupial antifeedant from Eucalyptus.

The resistance of Eucalyptus to browsing mammals has been related to the level and type of formylated phloroglucinol compounds (FPCs) present in the leaf. The antifeedant activity of FPCs appears to depend on their aldehyde groups, but little else is known of their mode of action. We have sought to elucidate this further by examining the biological reactivity and disposition of jensenone, a model FPC. Neither jensenone nor any metabolites were detected in urine or feces of marsupial brushtail or ringtail possums that had ingested up to 725 mg x kg(-0.75). When jensenone was incubated in rat gastrointestinal segments in vitro, it rapidly disappeared. Jensenone also reacted rapidly with glutathione, cysteine, glycine, ethanolamine, and trypsin, and more slowly with acetylcysteine and albumin. Sideroxylonal, a more complex FPC, exhibited the same reactivity. Torquatone, a related compound that lacks both aldehyde groups and antifeedant activity, was unreactive. Mass spectroscopic analysis indicated that the adducts were Schiff bases formed between the aldehyde groups of FPCs and amine groups of the conjugating molecules. Successive adducts were formed with the two aldehyde groups of jensenone, and the four groups of sideroxylonal. The jensenone bis-glutathione adduct appeared to cyclize to the disulfide form. These findings suggest that the antifeedant effects of FPCs are due to their facile binding to amine groups on critical molecules in the gastrointestinal tract, leading to a loss of metabolic function. The consequent toxic reaction, probably involving chemical mediators such as 5-hydroxytryptamine (5HT), may cause colic, nausea, and a general malaise, resulting in anorexia.

Adaptation, Physiological↗

Antiherbivore chemistry of Eucalyptus-cues and deterrents for marsupial folivores.

Formylated phloroglucinol compounds (FPCs) are the single most important factor determining the amount of foliage that marsupial folivores eat from individual Eucalyptus trees. Folivores need to recognize which trees contain FPCs if they are to avoid them and forage efficiently, they are challenged by great diversity in the types and quantities of FPCs present, even within eucalypt species. We investigated the relationship between FPCs and terpenoids in species with both simple and complex FPC profiles and found strong positive correlations between terpenes generally, and several monoterpenes in particular, and FPCs. Terpene cues also indicated qualitative differences in trees' FPC profiles. We describe significant qualitative and quantitative variation in FPCs in several species that are important food sources for marsupial folivores. New discoveries include the fact that macrocarpals occur as two major, distinct groups and several new dimeric acylphloroglucinols from Eucalyptus strzeleckii. These patterns add to the chemical complexity of the foraging environment for folivores.

Animals↗

Glucuronuria in the koala.

Glucuronuria is normal in marsupial folivores such as the koala (Phascolarrctos cinereus), which excretes 2-3 g glucuronic acid daily. Although this has long been attributed to the metabolites of Eucalyptus terpenes, we have found that these are mostly excreted in the unconjugated form. We now report on the aglycones that account for most of the glucuronic acid in koala urine. Urine (24 hr) was collected from six male koalas (8.8 +/- 0.4 kg, mean +/- SE) that were maintained on E. cephalocarpa foliage. Urine samples were analyzed by liquid and gas chromatography (LC and GC) coupled with mass spectrometry (MS). Glucuronides were readily identified by LC-MS/MS, which generated characteristic product ions at m/z 113 and 175. From the corresponding parent glucuronide ions, the masses of the aglycones were calculated. Confirmation of identity was by GC-MS after hydrolysis with beta-glucuronidase and comparison with standard compounds. Quantitation was by GC. The major non-terpene aglycones were 4-methylcatechol, resorcinol, salicyl alcohol, and two unidentified C7H8O2 phenols. Smaller amounts of benzoic acid, benzyl alcohol, orcinol, p-cresol, phenol, and phloroglucinol were detected. We have previously reported that terpene metabolites account for about 10% urinary glucuronides in the same koalas fed E. cephalocarpa. The present study found that an additional 60% urinary glucuronic acid is conjugated with non-terpene, mainly phenolic, aglycones. It seems likely that these phenolic compounds are present in leaves as glycosides and are chiefly responsible for the glucuronuria in koalas.

Animals↗

Quantification of sideroxylonals in Eucalyptus foliage by high-performance liquid chromatography.

This paper describes the extraction and quantification of sideroxylonals, a group of formylated phloroglucinol compounds found in the foliage of some eucalypt species. Samples of dry, ground foliage were Soxhlet-extracted with light petroleum spirit:acetone (4:1) and the resultant extract analysed (in the presence of internal standard) by reversed-phase HPLC without further purification. The yield of sideroxylonals was exponential with time and showed an inflection at ca. 4 h of extraction. It is recommended that samples be extracted for 6 h, giving a 92% recovery of the sideroxylonals. The title compounds deteriorate under various conditions, e.g. 10% are lost when foliage is oven-dried at 40 degrees C compared to freeze-drying. Storing samples in mobile phase led to a slow deterioration of sideroxylonals with a 7% loss after 4 days, while 22% of these compounds were lost from dry, ground eucalypt leaf stored at room temperature for 20 months.

Benzofurans↗

The rapid determination of sideroxylonals in Eucalyptus foliage by extraction with sonication followed by HPLC.

A rapid method is described for the quantification of sideroxylonals, a group of formylated phloroglucinol compounds found in some eucalypts. Samples of dry, ground foliage were extracted by sonication with 20% methanol in acetonitrile, 7% water in acetonitrile or 40% water in acetonitrile and the extracts analysed by reversed phase HPLC. The extracts from the two water-acetonitrile extractions were stable for at least 48 h. All three sonication methods recovered more sideroxylonals than did the Soxhlet extraction with petroleum spirit and acetone. Adding 0.1% trifluoracetic acid to the water-acetonitrile extraction solvents led to even higher recoveries of sideroxylonals. Soaking the sample in extracting solvent for 5 min recovered 70% of the sideroxylonals, whilst sonicating the suspension for 1 min recovered the remainder. The developed method involving sonication of the sample for 5 min in 7% water in acetonitrile with 0.1% trifluoroacetic acid is fast and requires minimal equipment and solvents compared with the traditional methods. With an autosampler it is possible to prepare and run 100 samples a day. More importantly, the technique is ideal for the analysis of small samples, e.g. individual leaves, which is essential when studying the evolutionary ecology of eucalypts.

Benzofurans↗