2.2. Characteristics
2.2.1 Importance
Because Azolla has a higher crude protein content (ranging from 19 to 30 percent) than most green forage crops and aquatic macrophytes and a rather favourable essential amino acid (EAA) composition for animal nutrition (rich in lysine), it has also attracted the attention of livestock, poultry and fish farmers (Cagauan and Pullin, 1991). In Asia Azolla has been long used as green manure for crop production and a supplement to diets for pigs and poultry. Some strains of Azolla can fix as much as 2-3 kg of nitrogen/ha/day.
Azolla doubles its biomass in 3-10 days, depending on conditions, and easily reaches a standing crop of 8-10 tonnes/ha fresh weight in Asian rice fields; 37.8 tonnes/ha fresh weight (2.78 tonnes/ha dry weight) has been reported for A. pinnata in India (Pullin and Almazan, 1983). Recently, Liu et al. (2008) have reported the application of Azolla as a controlled ecological life support system (CELSS) for its strong photosynthetic oxygen-releasing capacity. Azolla provides a protected environment and a fixed source of carbon to the blue-green filamentous algae Anabaena spp. (Wagner, 1997).
2.2.2 Environmental requirements
The potential for rearing Azolla is restricted by climatic factors, water and inoculum availability, the incidence of pests, phosphorus requirements and the need for labour intensive management (Cagauan and Pullin, 1991). Water is the fundamental requirement for the growth and multiplication of Azolla. The plant is extremely sensitive to lack of water. Although Azolla can grow on wet mud surfaces or wet pit litters, it prefers growing in a free-floating state (Becking, 1979). A strip of water not more than a few centimetres deep favours growth because it provides good mineral nutrition, with the roots not too far from the soil, and also because it reduces wind effects (Van Hove, 1989). Strong winds can accumulate Azolla to one side of the stretch of water, creating an overcrowded condition and thus slowing growth.
Azolla can survive a water pH ranging from 3.5-10, reported optimum growth occurring at pH 4.5-7.0. Watanabe et al. (1977) reported that the growth of Azolla was optimum at pH 5.5 and FAO (1977) recorded that soils of pH 6 to 7 support the best growth.
Salinity tolerance of Azolla species also varies. The growth rate of A. pinnata declines as its salinity increases above 380 mg/l (Thuyet and Tuan, 1973). According to Reddy et al. (2005) Azolla can withstand salinity of up to 10 ppt but Haller, Sutton and Burlowe (1974) reported that the growth of A. caroliniana ceases at about 1.3 ppt and higher concentrations result in death. A. filiculoides has been reported to be most salt-tolerant (I. Watanabe pers. comm., cited by Cagauan and Pullin, 1991).
Azolla grows in full to partial shade (100-50 percent sunlight) with growth decreasing quickly under heavy shade (Ferentinos et al., 2002). Generally, Azolla requires 25-50 percent full sunlight for its normal growth; slight shade is of benefit to its growth in field conditions. The biomass production greatly decreases at a light intensity lower than 1 500 lux (Liu et al., 2008).
Like all other plants, Azolla needs all the macro- and micro-nutrients for its normal growth and vegetative multiplication. All elements are essential; phosphorus is often the most limiting element for its growth. For normal growth, 0.06 mg/l/day is required. This level can be achieved in field conditions by the split application of superphosphate at 10-15 kg/ha (Sherief and James, 1994). 20 mg/l is the optimum concentration (Ferentinos et al., 2002). The symptoms of phosphorous deficiency are red-coloured fronds (due the presence of the pigment anthocyanin), decreased growth and curled roots. Macronutrients such as P, K, Ca and Mg and micronutrients such as Fe, Mo and Co have been shown to be essential for the growth and nitrogen fixation of Azolla (Khan and Haque, 1991).
The temperature tolerance of Azolla varies widely between its species and strains. In general, Azolla has low tolerance to high temperature and that restricts its use in tropical agriculture. There are, however, strains that can adapt successfully to high temperature. Cagauan and Pullin (1991) ranked different Azolla species from the most to the least heat-tolerant, based on the optimum temperature for good growth: A. mexicana > A. pinnata var. pinnata > A. microphylla > A. pinnata var. imbricata, A. caroliniana > A. filiculoides (Table 2.1). In general, the optimum temperature for growth of all Azolla species is around 25 ?C, except that of A. mexicana, whose optimum temperature is ~30 ?C.
Table 2.1
Temperature tolerance of five species of Azolla
According to Sherief and James (1994), the favourable water temperature for rapid multiplication of Azolla is generally between 18 and 26 ?C. The optimum relative humidity for Azolla growth is between 85-90 percent. Azolla becomes dry and fragile at a relative humidity lower than 60 percent.