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Two hypotheses, nutrient constraints and detoxification limitation, have been proposed to explain the lack of specialists among mammalian herbivores. The nutrient constraint hypothesis proposes that dietary specialization in mammalian herbivores is rare because no one plant can provide all requisite nutrients. The detoxification limitation hypothesis suggests that the mammalian detoxification system is incapable of detoxifying high doses of similar secondary compounds present in a diet of a single plant species. We experimentally tested these hypotheses by comparing the performance of specialist and generalist woodrats (Neotoma) on a variety of dietary challenges. Neotoma stephensi is a narrow dietary specialist with a single species, one-seeded juniper, Juniperus monosperma, comprising 85-95% of its diet. Compared with other plants available in the habitat, juniper is low in nitrogen and high in fiber, phenolics, and monoterpenes. The generalist woodrat, N. albigula, also consumes one-seeded juniper, but to a lesser degree. The nutrient constraint hypothesis was examined by feeding both species of woodrats a low-nitrogen, high-fiber diet similar to that found in juniper. We found no differences in body mass change, or apparent digestibility of dry matter or nitrogen between the two species of woodrats after 35 days on this diet. Moreover, both species were in positive nitrogen balance. We tested the detoxification limitation hypothesis by comparing the performance of the generalist and specialist on diets with and without juniper leaves, the preferred foliage of the specialist, as well as on diets with and without α-pinene, the predominant monoterpene in juniper. We found that on the juniper diet, compared with the specialist, the generalist consumed less juniper and lost more mass. Urine pH, a general indicator of overall detoxification processes, declined in both groups on the juniper diet. The generalist consumed half the toxin load of the specialist yet its urine pH was slightly lower. Moreover, the generalist consumed significantly less of the treatment with high concentrations of α-pinene compared to the control treatment, while the specialist consumed the same amount of food regardless of α-pinene concentration. For both groups, urine pH declined as levels of α-pinene in the diet increased. The generalist produced a significantly more acidic urine than the specialist on the treatment with the highest α-pinene concentration. Our results suggest that in this system, specialists detoxify plant secondary compounds differently than generalists and plant secondary compounds may be more important than low nutrient levels in maintaining dietary diversity in generalist herbivores.

Various plant secondary metabolites related to cinnamic acid are of interest because of their repellency to birds and their occurrence in ecologically important food items. Coniferyl benzoate (CB), a phenylpropanoid ester that occurs in quaking aspen (Populus tremuloides) is of particular ecological interest because of its effect on ruffed grouse (Bonasa umbellus) feeding behavior and its possible influence on the population dynamics of this bird. During detoxification processes, CB and other analogous compounds are metabolized into by-products, such as ferulic acid (FA), that can cause anti-reproductive effects. We tested whether consumption of CB produces antire-productive effects similar to FA using male and female Japanese quail (Coturnix coturnix) as avian models for ruffed grouse. The parameters we investigated included: the production, morphology, and development of eggs; reproductive characteristics influenced by estrogen; serum prolactin levels; and male reproductive behavior. Dietary CB did not produce antireproductive effects similar to FA at intake levels that Japanese quail and ruffed grouse would freely consume. Consumption of CB by Japanese quail significantly reduced egg production and body mass but did not affect male reproductive performance. Coniferyl benzoate's effect on egg production may be explained by lower energy acquisition and retention rather than endocrine changes per se. Contrary to previous reports, it is unlikely that FA, or similar compounds act directly as estrogen mimics or antagonists. Although, CB did reduce egg production in quail, it is unlikely that it would affect egg production in wild ruffed grouse. Detoxification costs and the effects of CB on nutrient utilization may explain why ruffed grouse avoid high dietary levels of CB.

The acute effects of tannin (tannic acid; TA) on nutrient absorption were studied by measuring sugar and amino acid uptake across the brash border (luminal membrane) of intact intestine in the presence and absence of TA. Incubation of tissue for 4-9 min in TA solution (1 mg/ml) caused a reduction in passive influx ofL-glucose in voles and mice and a reduction in carrier-mediated influx ofD-glucose and total influx ofL-proline in mice, but not voles. In subchronic experiments, mice and voles were fed for 7-14 days a diet with 4% TA, but there was no significant effect on intestinal brush border uptake ofL-glucose,D-glucose, orL-proline (or three other amino acids tested in voles). In a synthesis of our study with others in the literature, three inferences are made from the patterns of effects across solutes, time scales of exposure, and species. First, the transport inhibitory effects following acute exposure are probably mediated by two processes: increased resistance to passive flux across an effective unstirred layer juxtaposed to the brush border membrane, perhaps due to tannin-mucin binding, and reduced Na(+)-coupled nutrient uptake across the intestinal brush border. Second, there is a species sensitivity difference in TA's effect on the second process. Third, the negative effects observed at the acute time scale in vitro do not necessarily occur in animals eating TA subchronically because little TA reaches the luminal membrane, or if it does its effects are quickly reversed when the tissue is removed and washed with solution free of TA.

Daily energy metabolism and water flux were measured with doubly labeled water in the free-living insectivorous lizards Cnemidophorus tigris (mean body mass 15.7 g) and Callisaurus draconoides (8.6 g) in June 1979 in the Colorado Desert of California. C. tigris was an active forager; it spent 91% of its 5-h daily activity period in movement. C. draconoides was a sit-and-wait predator; it spent less than 2% of its 10-h activity period in movement. C. tigris had significantly higher rates of field energy metabolism and water influx (210 Jg -1 day -1 , 36.8 ul H 2 O g -1 day -1 , N=19) than C. draconoides (136, 17.1, N=18). There were no significant differences between the sexes within either species.The extra costs of free existence were calculated from differences between field metabolic rates and maintenance costs estimated from laboratory respirometry. Rates of energy metabolism during the field activity period were about 1.5x resting levels at 40° C (∼field active body temperature) for C. draconoides and 3.3 x resting levels at 40° C (∼field active body temperature) for the more active C. tigris. Feeding rates calculated from water influx data were 13.3 mg g -1 day -1 for C. tigris and 5.8 mg g -1 day -1 for C. draconoides. Though C. tigris had a high rate of energy expenditure, its foraging efficiency [Formula: see text] was higher than C. draconoides'.