The cassava plant, Manihot esculenta, grows exceptionally well in low fertility and drought prone environments, but the mechanisms that allow this growth are unknown. Earlier, and sometimes contradictory, work speculated about the presence of a C4-type photosynthesis in cassava leaves. In the present work we found no evidence for a C4 metabolism in mature attached cassava leaves as indicated i) by the low, 2 to 8%, incorporation of (14)CO2 into C4 organic acids in short time periods, 10 s, and the lack of (14)C transfer from C4 acids to other compounds in (12)CO2, ii) by the lack of C4 enzyme activity changes during leaf development and the inability to detect C4 acid decarboxylases, and iii) by leaf CO2 compensation values between 49 and 65 μl of CO2 1(-1) and by other infrared gas exchange photosynthetic measurements. It is concluded that the leaf biochemistry of cassava follows the C3 pathway of photosynthesis with no indication of a C3-C4 mechanism.However, cassava leaves exhibit several novel characteristics. Attached leaves have the ability to effectively partition carbon into sucrose with nearly 45% of the label in sucrose in about one min of (14)CO2 photosynthesis, contrasting with 34% in soybean (C3) and 25% in pigweed (C4). Cassava leaves displayed a strong preference for the synthesis of sucrose versus starch. Field grown cassava leaves exhibited high rates of photosynthesis and curvilinear responses to increasing sunlight irradiances with a tendency to saturate only at high irradiances, above 1500 μmol m(-2) s(-1). Morphologically, the cassava leaf has papillose epidermal cells on its lower mesophyll surface that form 'fence-like' arrangements encircling guard cells. It is proposed that the active synthesis of sugars has osmotic functions in the cassava plant and that the papillose epidermal cells function to maintain a healthy leaf water status in various environments.
About 95% of swamp tupelo (Nyssa sylvatica var. biflora (Walt.) Sarg.) and sweetgum (Liquidambar styraciflua L.) seedlings survived continuous root flooding for more than two years, whereas none of the swamp chestnut oak (Quercus michauxii Nutt.) and cherrybark oak (Q. falcata var. pagodifolia Ell.) seedlings survived one year of flooding. Death of oak seedlings occurred in phases associated with periods of major vegetative growth, e.g., after bud burst in spring, after summer stem elongation, and during the winter deciduous stage, suggesting that stored reserves and sources were inadequate to maintain the seedlings when vegetative sinks were forming. Additional evidence that flooding induced a source deficiency in oak was that leaves of flooded oak were 65 to 75% smaller than leaves of nonflooded oak. Flooded swamp tupelo seedlings had a normal leaf size and patchy stomatal opening compared with nonflooded seedlings. Flooding caused increases in alcohol dehydrogenase (ADH) specific activity in taproot cambial tissues and increases in starch concentrations of swamp tupelo seedlings that were reversed when seedlings were removed from flooding. Flooding had little effect on soluble sugar concentrations in swamp tupelo or sweetgum. In the long-term flood-dry-flood treatment, in which all species had survivors, upper canopy leaf photosynthetic rates were higher in all species during the dry period than in nonflooded controls, whereas their starch and soluble sugars concentrations were similar to those of nonflooded controls. Based on seedling survival and the sink-source relationships, the order of flood tolerance was: swamp tupelo > sweetgum > swamp chestnut oak > cherrybark oak.
The hypothesis we propose is that during photosynthesis the balance between potentially detrimental and beneficial photochemically induced events can be tipped beneficially toward increased photosynthesis and toward increased crop yield. To test this hypothesis a procedure has been devised with the rice plant, Oryza sativa, that has resulted in increasing both canopy photosynthesis and rice grain yield. Two elite rice varieties selected independently in the contrasting environments of either South China or Texas, each with distinct photosynthetic traits, were crossed to produce a hybrid with an increased canopy photosynthesis and grain yield that is regularly 20 to 22% higher than the mid-yields of the parents. The photosynthetic and mechanisms which may contribute to these beneficial results in the hybrid rice are: a reduction of the midday depression of photosynthesis; a rapid development of the canopy for photosynthetic light interception and an increased canopy photosynthesis; increased amounts of carotenoids for the xanthophyll cycle; an increased protection against free radicals induced by paraquat treatment; a 6 to 12 day shorter plant reproductive life cycle; and a 8 to 10 day increase in the longevity of the flag leaf over the parents. While the hybrid rice has successfully integrated these and likely other unknown characteristics to increase both crop photosynthesis and grain yield, we propose that understanding the underlying beneficial photosynthetic mechanisms supporting these crop plant traits is worthy of thorough investigation and application in crop production.
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