International Journal of Aquaculture, 2013, Vol.3, No.13, 63
          
        
        
          -
        
        
          
            72
          
        
        
        
          64
        
        
          Kohno et al., 1990; Kohno et al., 1997). Difficulties in
        
        
          rearing early stage larvae of groupers have become the
        
        
          major bottleneck hindering the development of mass
        
        
          fingerling production (Kohno et al., 1997). Marte (2003)
        
        
          summarized the difficulty of rearing grouper into three
        
        
          area: 1) spawned eggs and larvae are very small and
        
        
          the small mouth gape in early larvae limits the choice
        
        
          of initial live feed; 2) grouper are extremely sensitive
        
        
          to mechanical disturbance; 3) long duration of larval
        
        
          rearing (>60 days).
        
        
          In this review, we will use the life cycle of grouper
        
        
          larvae as a framework to review internal factors
        
        
          regulating ontogenetic development in fish larvae and
        
        
          environmental factors affecting the general develoment of
        
        
          grouper larvae. To understand the cause of high larvae
        
        
          mortality in early life history, we will review factors
        
        
          related to the ontogenetic development, and then we
        
        
          focus on issues of first feeding of grouper larvae in
        
        
          intensive aquaculture. At the end, we will review
        
        
          the management strategies of using live feeds in
        
        
          grouper hatcheries.
        
        
          
            1
          
        
        
          
            Ontogenetic Development
          
        
        
          
            1.1
          
        
        
          
            Eggs and Embryo
          
        
        
          The eggs size of groupers is generally less than one
        
        
          millimeter (Table 1). Like most marine teleosts,
        
        
          nutrition during the embryonic phase is derived from
        
        
          yolk reserve (Ma et al., 2012). The embryonic period
        
        
          starts from fertilization and ends at the commencement
        
        
          of exogenous feeding. It is divided into three major
        
        
          phase: cleavage egg, embryo, and free embryo (Moyle
        
        
          and Cech, 2003). Figure 1 illustrates the embryonic
        
        
          development of Malabar grouper
        
        
          
            Epinephelus
          
        
        
          
            malabaricus
          
        
        
          .
        
        
          Cleavage egg (A-F), and embryo (G-L)
        
        
          are defined according to Moyle and Cech (2003).
        
        
          Embryonic development is a complex process and egg
        
        
          quality and hatching environments directly affect
        
        
          embryonic development and the sizes of fish at
        
        
          hatching and first feeding (Robin and Gatesoupe, 2001).
        
        
          Table 1 Comparison of the eggs and larvae of
        
        
          
            Epinephaline
          
        
        
          Serranids
        
        
          Species
        
        
          Egg diameter
        
        
          (
        
        
          mm)
        
        
          Incubation temperature (
        
        
          ℃
        
        
          )
        
        
          Hatching
        
        
          time (h)
        
        
          Length at hatching
        
        
          (
        
        
          mm)
        
        
          Duration of Yolksac
        
        
          absorbation (day)
        
        
          Reference
        
        
          
            Epinephelus
          
        
        
          
            malabaricus
          
        
        
          0.87
        
        
          -
        
        
          0.93
        
        
          25
        
        
          -
        
        
          31
        
        
          29
        
        
          -
        
        
          26
        
        
          1.71
        
        
          -
        
        
          1.98
        
        
          3
        
        
          -
        
        
          4
        
        
          Leu et al., 2005;
        
        
          Yoseda et al., 2006
        
        
          
            E. tauvina
          
        
        
          0.80
        
        
          -
        
        
          0.90
        
        
          27
        
        
          -
        
        
          30
        
        
          25
        
        
          -
        
        
          20
        
        
          1.50
        
        
          -
        
        
          2.4
        
        
          3
        
        
          Lim 1993; Hussain
        
        
          et al., 1980
        
        
          
            E. fuscoguttatus
          
        
        
          0.89
        
        
          28
        
        
          -
        
        
          30
        
        
          19
        
        
          -
        
        
          18
        
        
          1.80
        
        
          -
        
        
          1.90
        
        
          -
        
        
          Lim 1993
        
        
          
            E. suillus
          
        
        
          28
        
        
          -
        
        
          30
        
        
          20
        
        
          -
        
        
          18
        
        
          1.62
        
        
          -
        
        
          Duray et al.,1996;
        
        
          Duray et al.,1997
        
        
          
            E. coioides
          
        
        
          0.74
        
        
          -
        
        
          0.85
        
        
          24.9
        
        
          -
        
        
          28
        
        
          21
        
        
          -
        
        
          19
        
        
          1.33
        
        
          -
        
        
          1.86
        
        
          3
        
        
          Doi et al.,1991;
        
        
          Zou et al., 2003
        
        
          
            E. costae
          
        
        
          0.89
        
        
          -
        
        
          0.95
        
        
          25
        
        
          -
        
        
          25.5
        
        
          28
        
        
          -
        
        
          24
        
        
          1.69
        
        
          -
        
        
          1.85
        
        
          3
        
        
          Glamuzina et al., 2000
        
        
          
            E. akaara
          
        
        
          0.70
        
        
          -
        
        
          0.77
        
        
          25
        
        
          -
        
        
          27
        
        
          25
        
        
          -
        
        
          23
        
        
          1.45
        
        
          -
        
        
          1.56
        
        
          4
        
        
          Ukawa et al., 1966
        
        
          
            E. polyphekadion
          
        
        
          0.71
        
        
          -
        
        
          0.83
        
        
          26
        
        
          -
        
        
          29
        
        
          21
        
        
          -
        
        
          19
        
        
          1.70
        
        
          -
        
        
          1.80
        
        
          2
        
        
          Rasem et al., 1997
        
        
          
            E. marginatus
          
        
        
          0.74
        
        
          -
        
        
          0.94
        
        
          23
        
        
          33
        
        
          -
        
        
          30
        
        
          1.40
        
        
          -
        
        
          1.67
        
        
          3
        
        
          -
        
        
          4
        
        
          Glamuzina et al., 1998
        
        
          
            E. merra
          
        
        
          0.71
        
        
          -
        
        
          0.73
        
        
          26.5
        
        
          -
        
        
          28.3
        
        
          27
        
        
          -
        
        
          24
        
        
          1.40
        
        
          -
        
        
          1.60
        
        
          2
        
        
          Jagadis et al., 2006
        
        
          
            1.1.1
          
        
        
          
            Egg quality
          
        
        
          Egg quality is generally derived from broodstock
        
        
          nutrition (Izquierdo et al., 2001; Mazorra et al., 2003;
        
        
          Sawanboonchun et al., 2008; Ma et al., 2012). Since
        
        
          protein, lipoprotein, glycogen, and enzymes contents
        
        
          in yolk reserve directly affect embryonic development
        
        
          (
        
        
          Gunasekera et al., 1995; Harrell and Woods, 1995;
        
        
          Sargent et al., 1999), proper controlled broodstocks
        
        
          nutrition is essential in breeding marine fish. For
        
        
          instance, Dhert et al. (1991) showed that
        
        
          
            E. tauvina
          
        
        
          broodstock given trash fish injected the emulsified
        
        
          enrichment diet Marila diet significantly increased oil
        
        
          globule diameter, total lipids, eicosapentaenoic acid,
        
        
          docosahexaenoic acid, and larval survival at day 7.
        
        
          Nutrients such as essential fatty acids (EFA)
        
        
          (
        
        
          Fernández-Palacios et al., 1995), vitamin E
        
        
          (
        
        
          Fernández-Palacios et al., 1998), carotenoids (Craik, 1985),
        
        
          vitamin C (Blom and Dabrowski, 1995), dietary
        
        
          protein, vitamin B
        
        
          1
        
        
          ,
        
        
          and vitamin B
        
        
          6
        
        
          (
        
        
          Izquierdo et al., 2001)
        
        
          in broodstock diets have been considered as essential