13.4 Physiological Rhythms: Matching Fish and Plant Nutrition
The design of feeds for fish is crucial in aquaponics because fish feed is the single or at least the main input of nutrients for both animals (macronutrients) and plants (minerals) (Fig. 13.3).
Nitrogen is introduced to the aquaponic system through protein in fish feed which is metabolized by fish and excreted in the form of ammonia. The integration of recirculating aquaculture with hydroponics can reduce the discharge of unwanted nutrients to the environment as well as generate profits. In an early economic study, phosphorus removal in an integrated trout and lettuce/basil aquaponic system proved to be cost-saving (Adler et al. 2000). Integrating fish feeding rates is also paramount to fulfil the nutritional requirements of plants. Actually, farmers need to know the amount of feed used in the aquaculture unit to calculate how much nutrient needs to be supplemented to promote plant growth in the hydroponic unit. For instance, in a tilapia-strawberry aquaponic system, the total amount of feed required to produce ions (e.g. NOsub3/subsup—,/sup Casub2/subsup+/sup, Hsub2/subPOsub4/subsup—/sup and Ksup+/sup) for plants was calculated at different fish densities, with better result for small fish density 2 kg fish/msup-3/sup to reduce the cost of hydroponic solution supplementation (Villarroel et al. 2011).
Fig. 13.3 Nutrient flow in an aquaponic system. Note that fish feed, through wastewater from the aquaculture system, provides the minerals required for plants to grow in the hydroponic system. Meal timing should be designed to match feeding/ excretion rhythms in fish and nutrient uptake rhythms in plants
It is well known that plants have daily rhythms and circadian rhythmicity in leaf movements was first described in plants by de Mairan in the early eighteenth century (McClung 2006). Circadian rhythms in plants control everything from the time of flowering to plant nutrition and thus these rhythms need to be taken into account especially when using artificial horticultural lighting. Fish are also tied into daily rhythms in most physiological functions, including feeding and nutrient uptake. It should not be surprising that fish exhibit feeding rhythms because food availability and the occurrence of predators are hardly constant but restricted to a particular time of day/night (López-Olmeda and Sánchez-Vázquez 2010). Thus, fish should be fed at the right time according to their appetite rhythms: meals scheduled during daytime for diurnal fish species, and at night for nocturnal fish. It is well known that fish show daily patterns of deamination of proteins and nitrogenous wastes related to their nutritional status and feeding rhythms (Kaushik 1980). Feeding time affects nitrogen excretion, as Gelineau et al. (1998) reported ammonia production and protein catabolism were lower in fish fed at dawn (in phase with their feeding rhythm) than in those fed at midnight (out of phase). Most interestingly, urea excretion shows circadian rhythmicity that persist in starved fish under constant conditions (Kajimura et al. 2002), revealing its endogenous origin. Furthermore, urea permeability (determined as body urea contents after immersion in a urea solution) coincided with the acrophase, i.e. the peak of the daily excretion rhythm, indicating that urea does not permeate cells by simple diffusion but there is a circadian control. Plants also show daily rhythms in nitrogen uptake, as early described by Pearson and Steer (1977), who found a daily pattern of nitrate uptake and nitrate reductase in peppers kept in a constant environment. The nitrate concentration in the leaves of spinach also increased during the night as the uptake rate of nitrate by the roots increase at that time (Steingrover et al. 1986). In aquaponics, the evidence thus points to the need for matching excretion rhythms in fish and nutrient uptake rhythms in plants. To optimize the performance and cost-effectiveness of aquaponic systems, fish diets and feeding schedules should be designed carefully to provide nutrients at the right level and the right time to complement both fish and plants.