Increased photosynthesis and water potentials in Silphium integrifolium galled by cynipid wasps

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Abstract
Interactions between drought, insect herbivory, photosynthesis, and water potential play a key role in determining how plants tolerate and defend against herbivory, yet the effects of insect herbivores on photosynthesis and water potential are seldom assessed. We present evidence that cynipid wasp galls formed by Antistrophus silphii on Silphium integrifolium increase photosynthesis (A), stomatal conductance (g), and xylem water potential (Ψ). Preliminary data showed that in drought-stressed plants galled shoots had 36% greater A, and 10% greater stem Ψ than ungalled shoots, while in well-watered plants leaf gas exchange was not affected by galls. We hypothesized that 1) galled shoots have higher Ψ, g, and A than ungalled shoots, but this differences diminishes if plant drought stress is reduced, and 2) galls can reduce decreases in A and g if water availability decreases. A field experiment testing the first hypothesis found that galls increased g and Ψ, but that differences between galled and ungalled shoots did not diminish after plants were heavily watered. A laboratory test of the second hypothesis using potted Silphium found that galled plants had smaller drops in A and g over a 4-day dry-down period. A vs g and A vs intercellular CO2 concentration relationships were consistent with the explanation that increased Ψ allows galls to increase A by reducing stomatal limitation of A, rather than by altering sink-source relationships or by removing low-Ψ limitations on non-stomatal components of A. Our working hypothesis is that galls increase Ψ and A by reducing the shoot: root ratio so that the plant is exploiting a greater soil volume per unit leaf area. We argue that increased A is an ineffective way for Silphium to compensate for negative effects of gall insect attack. Instead, increased Ψ and A may protect gall insects from variation in resource availability caused by periodic drought stress, potentially reducing negative effects of drought on plant quality and on gall insect populations.