Previous work has shown that stomata respond directly to light, but it was not clear whether the only additional response is through CO2, or whether some other metabolite is involved in this response. Gas exchange experiments were done with normally positioned and inverted leaves of Hedera helix to investigate this problem. The macroscopic optical properties of the leaves and their anatomical structure were also studied. These experiments snowed that there is no need to postulate the existence of a messenger other than CO2 to explain the indirect response of stomata to light. The experiments also showed that leaf inversion affects both stomatal conductance and photosynthesis, and highlight the difficulties involved in the interpretation of the effect of leaf inversion on stomata when stomatal conductance measurements are not done concurrently with measurements of CO2 flux density and intercellular CO2 molar fraction
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Aphalo et al. (1993) studied this question.