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Cassava or manioc (Manihot esculenta Crantz), a perennial shrub of the New World, currently is the sixth world food crop for more than 500 million people in tropical and sub-tropical Africa, Asia and Latin America. It is cultivated mainly by resource-limited small farmers for its starchy roots, which are used as human food either fresh when low in cyanogens or in many processed forms and products, mostly starch, flour, and for animal feed. Because of its inherent tolerance to stressful environments, where other food crops would fail, it is often considered a food-security source against famine, requiring minimal care. Under optimal environmental conditions, it compares favorably in production of energy with most other major staple food crops due to its high yield potential. Recent research at the Centro Internacional de Agricultura Tropical (CIAT) in Colombia has demonstrated the ability of cassava to assimilate carbon at very high rates under high levels of humidity, temperature and solar radiation,which correlates with productivity across all environments whether dry or humid. When grown on very poor soils under prolonged drought for more than 6 months, the crop reduce both its leaf canopy and transpiration water loss, but its attached leaves remain photosynthetically active, though at greatly reduced rates. The main physiological mechanism underlying such a remarkable tolerance to drought was rapid stomatal closure under both atmospheric and edaphic water stress, protecting the leaf against dehydration while the plant depletes available soil water slowly during long dry periods. This drought tolerance mechanism leads to high crop water use efficiency values. Although the cassava fine root system is sparse, compared to other crops, it can penetrate below 2 m soil,thus enabling the crop to exploit deep water if available. Leaves of cassava and wild Manihot possess elevated activities of the C4 enzyme PEP carboxylase but lack the leaf Kranz anatomy typical of C4 species, pointing to the need for further research on cultivated and wild Manihot to further improve its photosynthetic potential and yield,particularly under stressful environments. Moreover, a wide range in values of Km (CO2) for the C3 photosynthetic enzyme Rubisco was found among cassava cultivars indicating the possibility of selection for higher affinity to CO2, and consequently higher leaf photosynthesis. Several plant traits that may be of value in crop breeding and improvement have been identified, such as an extensive fine root system, long leaf life, strong root sink and high leaf photosynthesis. Selection of parental materials for tolerance to drought and infertile soils under representative field conditions have resulted in developing improved cultivars that have high yields in favorable environments while producing reasonable and stable yields under stress.

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The paper summarizes research conducted at International Center for Tropical Agriculture (CIAT) on responses of cassava to extended water shortages in the field aided by modern gas-exchange and water-relation techniques as well as biochemical assays. The aim of the research was to coordinate basic and applied aspects of crop physiology into a breeding strategy with a multidisciplinary approach. Several physiological characteristics/traits and mechanisms underpinning tolerance of cassava to drought were elucidated using a large number of genotypes from the CIAT core germplasm collection grown in various locations representing ecozones where cassava is cultivated. Most notable among these characteristics are the high photosynthetic capacity of cassava leaves in favorable environments and the maintenance of reasonable rates throughout prolonged water deficits, a crucial characteristic for high and sustainable productivity. Cassava possess a tight stomatal control over leaf gas exchange that reduces water losses when plants are subjected to soil water deficits as well as to high atmospheric evaporative demands, thus protecting leaves from severe dehydration. During prolonged water deficits, cassava reduces its canopy by shedding older leaves and forming smaller new leaves leading to less light interception, another adaptive trait to drought. Though root yield is reduced (but much less than the reduction in top growth) under water stress, the crop can recover when water becomes available by rapidly forming new canopy leaves with much higher photosynthetic rates compared to unstressed crops, thus compensating for yield losses with final yields approaching those in well-watered crops. Cassava can extract slowly water from deep soils, a characteristic of paramount importance in seasonally dry and semiarid environments where deeply stored water needs to be tapped. Screening large accessions under seasonally dry and semiarid environments showed that yield is significantly correlated with upper canopy leaf photosynthetic rates, and the association was attributed mainly to nonstomatal (anatomical/biochemical) factors. Parental materials with both high yields and photosynthetic rates were identified for incorporation into breeding and selection programs for cultivars adapted to prolonged drought coupled with high temperatures and dry air, conditions that might be further aggravated by global climate changes in tropical regions. Key words: agriculture, breeding, canopy, carboxylation enzymes, C 3 -C 4 intermediate, ecophysiology, growth, leaf photosynthesis, Manihot sp., PEP carboxylase, productivity, stomata, water stress, yield Características fisiológicas da tolerância da mandioca a secas prolongadas nos trópicos: Implicações para o melhoramento de cultivares adaptadas a ambientes semi-áridos e sazonalmente secos: Este artigo sumaria resultados de pesquisas conduzidas no Centro Internacional para Agricultura Tropical (CIAT), sobre as respostas da mandioca a períodos prolongados de déficit hídrico, no campo, centrando-se em...

exchange and stomatal/non-stomatal limitation to CO 2 assimilation of Quercus liaotungensis under midday high irradiance.-Photosynthetica 39: 383-388, 2001. ERRATUM M.A. El-Sharkawy: International research on cassava photosynthesis, productivity, eco-physiology, and responses to environmental stresses in the tropics. − Photosynthetica 44: 481-512, 2006.

The review sums up research conducted at CIAT within a multidiscipline effort revolving around a strategy for developing improved technologies to increase and sustain cassava productivity, as well as conserving natural resources in the various eco-edaphic zones where the crop is grown, with emphasis on stressful environments. Field research has elucidated several physiological plant mechanisms underlying potentially high productivity under favourable hot-humid environments in the tropics. Most notable is cassava inherent high capacity to assimilate carbon in near optimum environments that correlates with both biological productivity and root yield across a wide range of germplasm grown in diverse environments. Cassava leaves possess elevated activities of the C 4 phosphoenolpyruvate carboxylase (PEPC) that also correlate with leaf net photosynthetic rate (P N ) in field-grown plants, indicating the importance of selection for high P N . Under certain conditions such leaves exhibit an interesting photosynthetic C 3 -C 4 intermediate behaviour which may have important implications in future selection efforts. In addition to leaf P N , yield is correlated with seasonal mean leaf area index (i.e. leaf area duration, LAD). Under prolonged water shortages in seasonally dry and semiarid zones, the crop, once established, tolerates stress and produces reasonably well compared to other food crops (e.g. in semiarid environments with less than 700 mm of annual rain, improved cultivars can yield over 3 t ha −1 oven-dried storage roots). The underlying mechanisms for such tolerance include stomatal sensitivity to atmospheric and edaphic water deficits, coupled with deep rooting capacities that prevent severe leaf dehydration, i.e. stress avoidance mechanisms, and reduced leaf canopy with reasonable photosynthesis over the leaf life span. Another stress-mitigating plant trait is the capacity to recover from stress, once water is available, by forming new leaves with even higher P N , compared to those in nonstressed crops. Under extended stress, reductions are larger in shoot biomass than in storage root, resulting in higher harvest indices. Cassava conserves water by slowly depleting available water from deep soil layers, leading to higher seasonal crop water-use and nutrient-use efficiencies. In dry environments LAD and resistance to pests and diseases are critical for sustainable yields. In semiarid zones the crop survives but requires a second wet cycle to achieve high yields and high dry matter contents in storage roots. Selection and breeding for early bulking and for medium/short-stemmed cultivars is advantageous under semiarid conditions. When grown in cooler zones such as in tropical high altitudes and in low-land sub-tropics, leaf P N is greatly reduced and growth is slower. Thus, the crop requires longer period for a reasonable productivity. There is a need to select and breed for more cold-tolerant genotypes. Selection of parental materials for tolerance to water stress and infertile soils has resulted in breeding ...

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SUMMARYYield stability in cassava requires genotypes that produce well under the variable moisture conditions encountered during the growth cycle. Plant characteristics related to yield stability were studied in two cassava clones subjected to 105 days of water stress in a field drainage lysimeter. Stress conditions commenced 117 days after planting, and the plants were allowed to recover at the end of the stress period for the rest of the growth cycle. Water stress restricted the growth of leaves and stems, but root yields were increased or remained unaffected. Leaf water potential varied little with stress, but gas exchange rates were about 75% those of the control throughout the stress period in both cultivars. Under stress, the plants partially closed their stomata and extracted deep soil moisture slowly. A high yield in both wet and stressed environments was associated with high mean LAI, better leaf retention, and greater partitioning of shoot biomass into leaf formation.