The concept that photosynthetic flux is influenced by the accumulation of p
hoto-assimilate persisted for 100 years before receiving any strong experim
ental support. Precise analysis of the mechanisms of photosynthetic respons
es to sink activity required the development of a battery of appropriate mo
lecular techniques and has benefited from contemporary interest in the effe
cts of elevated CO2 on photosynthesis. Photosynthesis is one of the most hi
ghly integrated and regulated metabolic processes to maximize the use of av
ailable light, to minimize the damaging effects of excess light and to opti
mize the use of limiting carbon and nitrogen resources. Hypotheses of feedb
ack. regulation must take account of this integration. In the short term, d
eparture from homeostasis can lead to redox signals, which cause rapid chan
ges in the transcription of genes encoding photosystems I and II. End-produ
ct synthesis can exert short-term metabolic feedback control through Pi rec
ycling. Beyond this, carbohydrate accumulation in leaves when there is an i
mbalance between source and sink at the whole plant level can lead to decre
ased expression of photosynthetic genes and accelerated leaf senescence. In
a high CO2 world this may become a more prevalent feature of photosyntheti
c regulation. However, sink regulation of photosynthesis is highly dependen
t on the physiology of the rest of the plant. This physiological state regu
lates photosynthesis through signal transduction pathways that co-ordinate
the plant carbon: nitrogen balance, which match photosynthetic capacity to
growth and storage capacity and underpin and can override the direct short-
term controls of photosynthesis by light and CO2. Photosynthate supply and
phytohormones, particularly cytokinins, interact with nitrogen supply to co
ntrol the expression of photosynthesis genes, the development of leaves and
the whole plant nitrogen distribution, which provides the dominant basis f
or sink regulation of photosynthesis.