Allocation of carbon to chemical defences has often been suggested to be a direct response to browsing or grazing by herbivores. This study quantitatively compares total polyphenol and tannin production in response to simulated browsing of three karoo shrubs in order to test this induced defence hypothesis. The three species studied responded to browsing either by rapid regrowth or by increasing polyphenol production in the remaining tissues. The patterns did not follow any phylogenetic relationships but were weakly associated with the palatability of each species. The highly palatable deciduous species Osteospermum sinuatum, which is capable of rapid regrowth, showed no or very low levels of constitutive and browsing-induced total polyphenols, condensed tannins and protein-precipitating tannins. The evergreen sclerophyllous species Pteronia pallens showed a limited regrowth capacity and had intermediate levels of polyphenols, while the evergreen succulent species Ruschia spinosa showed almost no regrowth over the study period. R. spinosa contained the highest constitutive and browsing-induced levels of polyphenols, condensed tannins and protein-precipitating tannins. In two of the species more than one anti-herbivore defence feature co-occur. P. pallens foliage contains both hepatotoxins and polyphenols while R. spinosa has both structural (spines) and chemical defences. Responses of karoo shrubs to simulated browsing are interpreted as the result of passive alterations in plant chemistry rather than as an active defence response to herbivores.
Nonstructural carbohydrate allocation patterns in response to different frequencies of simulated browsing (leaf and twig removal) were studied in the following semi-arid shrubs: Osteospermum sinuatum, a dwarf deciduous shrub, Pteronia pallens, a dwarf evergreen shrub, and Ruschia spinosa, a dwarf leaf-succulent shrub. Simulated browsing at all frequencies resulted in the elevation, or had no effect, on total nonstructural carbohydrate (TNC) concentrations of O. sinuatum plant parts, and resulted in the decrease in TNC concentrations of R. spinosa plant parts. The responses of P. pallens were intermediate with elevations as well as declines in TNC concentrations of plant parts measured in response to various clipping frequencies. At the low frequency of simulated browsing (every 26 weeks) elevations in plant TNC content were measured in the two non-succulent shrubs O. sinuatum and P. pallens. It was concluded that the overcompensation with respect to TNC accumulation observed in the two non-succulent species represents one of the ways in which excess photosynthate is utilized by browsed shrubs with a limited regiowth potential. Simulated browsing was the least detrimental with respect to biomass production to the non-succulent O. sinuatum and P. pallens, and most injurious to the leaf-succulent shrub, R. spinosa. The observed TNC allocation patterns could not adequately explain the variation among species in the production of new growth and it was concluded that some factor(s) other than the carbon resource was limiting regrowth.
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