Virtually all studies of effect of aphids on plant growth involve cultivated varieties of plants growing in unnatural assemblages. Choudhury's report is no exception as he chooses cultivated pea, Pisum sativum, which also happens to fix its own nitrogen and is therefore unlikely to benefit from a sugar input which, according to our hypothesis, stimulates nitrogen fixation by free-living soil bacteria. Choudhury's experiment is a good one in that he measures production of flowers and seeds rather than production of leaves, but like others (e.g. Dyer 1980, Petelle 1980) he uses wrong plant in wrong situation. What is required is an analysis of effect of a natural plant-feeder on a naturally growing plant. Only in this situation will it be possible to demonstrate presence or absence of mutualistic co-evolution. What then, would be a good choice? One is wild cabbage, Brassica oleracea, a relatively long-lived perennial confined to sea coasts of western Europe. In marked contrast to its many annual or biennial cultivars that are often severely damaged when grown under unnatural conditions, wild cabbage shows little evidence of detrimental effects of consumers, even though all cabbage-feeding specialists, together with an array of generalist feeders, are present (Owen unpubl.). As we previously pointed out (Owen and Wiegert 1976), many species of aphids aggregate and drain nutrients from distant parts of plants and in effect act as sinks, competing with natural sinks such as growing parts and storage organs. Even cultivars growing in unnatural conditions may occasionally 'benefit' from feeding by aphids, when 'benefit' is defined in terms of a product useful to man. Way and Cammell (1970), working with cabbage aphid, Brevicoryne brassicae, have shown that in cabbage supply of photosynthetic material exceeds capacity of plant's sink and so sugar accumulates in leaves and slows down photosynthesis which, in turn, decreases productivity. But our original, and continuing position has been that 'benefit' must be defined solely in terms of relative fitness, which is not necessarily related to productivity. Excess sugars released by aphids and deposited below plant could stimulate nitrogen fixation which would be beneficial for a plant unable to fix its own nitrogen and which very often grows in nitrogen deficient soils. We therefore agree with Choudhury that long-lived perennials are more likely to enter into a mutualistic relationship with consumers than short-lived annuals. Indeed we have never agreed with Stenseth's (1978) reasoning that short-lived r-strategists are more likely to benefit than long-lived K-strategists. Stenseth's conclusion stems directly from Leslie matrix model used in his theoretical analysis in which there is no threshold level below which grazing does not affect mortality. Using such a model, grazing can confer benefits to fitness (measured as rmax in Stenseth's paper) only when increased natality more than counteracts increased mortality due to grazing, in Stenseth's words the relative gain... must be greater longer life cycle. But we argue that grazing (or browsing), if regulated below given levels (which would be species specific), seldom if ever increases mortality of long-lived perennials. Thus, increased fitness of these organisms depends entirely on whether grazing produces more propagules with increased relative survival rates. This effect, we argue, is much more likely to be selected for when organism being grazed is long-lived, remaining in place to reap benefit of (in aphid example) whatever increased concentrations of nutrients are produced by action of aphids. Lastly, Choudhury seems to want to join increasing band of ecologists who see consumers as playing roles in ecosystem regulation. Although it may be true that there is some ecosystem effect, we maintain that evolutionary significance of plant-consumer relationships must be sought in terms of benefit, or otherwise, at individual plant level. Any regulatory effect on ecosystem is a by-product of organismorganism interactions whether they are mutualisms or not.
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