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Crop yield response to water - Cra

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with typical values around 10.5 g/m 2 under the low, natural fertility in the Bolivian Altiplano(Geerts et al., 2009). Still-under poor fertility conditions, a decrease of the reference biomass<strong>water</strong> productivity (WP*) value only occurs at higher <strong>to</strong>tal transpiration sums, and only by10 percent. The C 3 pathway is well adapted <strong>to</strong> the prevailing low average temperatures in theAltiplano. Reported values of seed <strong>yield</strong> per unit of <strong>water</strong> consumed (WP y/ET ) are rather low andlie between 0.3 and 0.6 kg/m³ because of the generally prevailing low fertility conditions (Geertset al., 2009). On the other hand, it is a crop with a large nitrogen-sink thus causing an increasedmetabolic cost or higher glucose-equivalent per unit dry matter produced. To our knowledge,no research has been conducted on the <strong>response</strong> of quinoa <strong>to</strong> increases in atmospheric CO 2 .Response <strong>to</strong> StressesQuinoa is highly resistant <strong>to</strong> a number of abiotic stresses (Jacobsen et al., 2003). Severaldrought resistant mechanisms are present in quinoa. Drought in early vegetative stages mayprolong its life cycle, allowing the plant <strong>to</strong> make up for growth lost during the early droughtif <strong>water</strong> is available later. Also, the availability of cultivars with different season length makesit possible <strong>to</strong> match the <strong>water</strong> requirement of the quinoa crop <strong>to</strong> the available rainfall or thes<strong>to</strong>red soil <strong>water</strong> at a given location. Quinoa tissue is relatively high in osmotic solutes andundergoessubstantial osmotic adjustment under drought, which enable s<strong>to</strong>mata <strong>to</strong> remainsomewha<strong>to</strong>pen down <strong>to</strong> a leaf <strong>water</strong> potential range of -1.5 MPa. During soil drying the plantsare able <strong>to</strong> maintain leaf <strong>water</strong> potential and pho<strong>to</strong>synthesis due <strong>to</strong> the complex s<strong>to</strong>matal<strong>response</strong>, resulting in an increase of leaf <strong>water</strong> use efficiency. Root originated ABA playsa role in s<strong>to</strong>mata performance during soil drying. ABA regulation seems <strong>to</strong> be one of themechanisms utilized by quinoa when facing drought inducing decrease of turgor of s<strong>to</strong>mataguard cells (Jacobsen et al., 2009). The plant also avoids negative effects of drought throughfast and deep rooting particularly in dry soils. Quinoa also reduces its leaf area by controlledleaf senescense under drought.Quinoa is a facultative halophyte (Bosque-Sanchez et al., 2003) and can grow in non-saline<strong>to</strong> extremely saline conditions, depending on the cultivar. Seed production is enhanced bymoderate salinity (EC in the 5-15 dS/m range) and may not be drastically reduced even at EC of40 <strong>to</strong> 50 dS/m in some cultivars (Jacobsen et al., 2003). Osmotic adjustment by the accumulationof salt ions in tissues enables the plant <strong>to</strong> maintain cell turgor and transpiration under salineconditions.Apart from drought, frost and cold are the other major growth limiting fac<strong>to</strong>rs in the Altiplano.Quinoa is <strong>to</strong>lerant <strong>to</strong> frost, partly because of the protection provided by its heterogeneouscanopy (Winkel et al., 2009), although the <strong>to</strong>lerance varies with cultivar and appears <strong>to</strong>diminish at the late phenological stages (Jacobsen et al., 2005). Leaf freezing of quinoaoccurred only between -5 and -6 °C, and is delayed in case of mild <strong>water</strong> stress (Bois et al.,2006). The resistance <strong>to</strong> frost is associated with super cooling of tissue <strong>water</strong> and <strong>to</strong>lerance ofextracellular ice formation (in the cell wall), as is common for most winter crops.Linked <strong>to</strong> frost resistance is a low base temperature (T b ase ) for plant processes. In a studyof the leaf appearance rate of different quinoa cultivars originating from various altitudesand latitudes, T base averaged 2 ºC and the temperature at which maximum rate was reachedaveraged 22 o C. Other studies found a T base of 3 °C, T opt of 30-35 °C and T max estimated <strong>to</strong> 50 °CQuinoa 233

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