Can fast-growing plantation trees escape biochemical down-regulation of photosynthesis when grown throughout their complete production cycle in the open air under elevated carbon dioxide?

Poplar trees sustain close to the predicted increase in leaf photosynthesis when grown under long-term elevated CO2 concentration ([CO2]). To investigate the mechanisms underlying this response, carbohydrate accumulation and protein expression were determined over four seasons of growth. No increase in the levels of soluble carbohydrates was observed in the young expanding or mature sun leaves of the three poplar genotypes during this period. However, substantial increases in starch levels were observed in the mature leaves of all three poplar genotypes grown in elevated [CO2]. Despite the very high starch levels, no changes in the expression of photosynthetic Calvin cycle proteins, or in the starch biosynthetic enzyme ADP-glucose pyrophosphorylase (AGPase), were observed. This suggested that no long-term photosynthetic acclimation to CO2 occurred in these plants. Our data indicate that poplar trees are able to 'escape' from long-term, acclimatory down-regulation of photosynthesis through a high capacity for starch synthesis and carbon export. These findings show that these poplar genotypes are well suited to the elevated [CO2] conditions forecast for the middle of this century and may be particularly suited for planting for the long-term carbon sequestration into wood.

Small decreases in SBPase cause a linear decline in the apparent RuBP regeneration rate, but do not affect Rubisco carboxylation capacity.

The response of net photosynthetic CO(2) uptake (A) to increasing leaf intercellular CO(2) concentration (c(i)) was determined in antisense Nicotiana tabacum plants, derived from six independent transformation lines, displaying a range of sedoheptulose-1, 7-bisphosphatase (SBPase) activities. The maximum in vivo ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) carboxylation (V(c,max)) and RuBP regeneration (J(max)) rates were calculated from the steady-state measurements of the A to c(i) response curves. In plants with reductions in SBPase activity of between 9% and 60%, maximum RuBP regeneration capacity declined linearly (r(2)=0.79) and no significant change in apparent in vivo Rubisco activity (V(c,max)) was observed in these plants. No correlation between V(c,max) and a decrease in capacity for RuBP regeneration was observed (r(2)=0.14) in the SBPase antisense plants. These data demonstrate that small decreases in SBPase activity limit photosynthetic carbon assimilation by reducing the capacity for RuBP regeneration.

Computer modelling and experimental evidence for two steady states in the photosynthetic Calvin cycle.

We present observations of photosynthetic carbon dioxide assimilation, and leaf starch content from genetically modified tobacco (Nicotiana tabacum) plants in which the activity of the Calvin cycle enzyme, sedoheptulose-1,7-bisphosphatase, is reduced by an antisense construct. The measurements were made on leaves of varying ages and used to calculate the flux control coefficients of sedoheptulose-1,7-bisphosphatase over photosynthetic assimilation and starch synthesis. These calculations suggest that control coefficients for both are negative in young leaves, and positive in mature leaves. This behaviour is compared to control coefficients obtained from a detailed computer model of the Calvin cycle. The comparison demonstrates that the experimental observations are consistent with bistable behaviour exhibited by the model, and provides the first experimental evidence that such behaviour in the Calvin cycle occurs in vivo as well as in silico.

Photosynthetic capacity is differentially affected by reductions in sedoheptulose-1,7-bisphosphatase activity during leaf development in transgenic tobacco plants.

The impact of reduced sedoheptulose-1,7-bisphosphatase (SBPase) activity on photosynthetic capacity and carbohydrate status was examined during leaf expansion and maturation in antisense transgenic tobacco (Nicotiana tabacum L. cv Samsun) plants. In wild-type plants, photosynthetic capacity was lowest in young expanding leaves and reached a maximum in the fully expanded, mature leaves. In contrast, the transgenic antisense SBPase plants had the highest photosynthetic rates in the young expanding leaves and lowest rates in the mature leaves. In the mature, fully expanded leaves of the transgenic plants photosynthetic capacity was closely correlated with the level of SBPase activity. However, in the youngest leaves of the SBPase antisense plants, photosynthetic rates were close to, or higher than, those observed in wild-type plants, despite having a lower SBPase activity than the equivalent wild-type leaves. Reductions in SBPase activity affected carbohydrate levels in both the mature and young developing leaves. The overall trend was for decreased SBPase activity to lead to reductions in carbohydrate levels, particularly in starch. However, these changes in carbohydrate content were also dependent on the developmental status of the leaf. For example, in young expanding leaves of plants with the smallest reductions in SBPase activity, the levels of starch were higher than in wild-type plants. These data suggest that the source status of the mature leaves is an important determinant of photosynthetic development.