|Arachidonic acid is a Major Component in Gonadal Fatty acids of Tropical
Coral Reef Fish in the Philippines and Japan
|Ashraf Suloma and Hiroshi Y. Ogata*
|Fisheries Division, Japan International Research Center for Agricultural Sciences, Ohwashi 1-1, Tsukuba 305-8686, Japan
||Dr. Hiroshi Y. Ogata
National Research Institute of
FRA, Tamaki, Mie 519-0423, Japan
|Received February 14, 2011; Accepted May 04, 2011; Published May 09 2011
|Citation: Suloma A, Ogata HY (2011) Arachidonic acid is a Major Component in
Gonadal Fatty acids of Tropical Coral Reef Fish in the Philippines and Japan. J
Aquac Res Development 2:111. doi:10.4172/2155-9546.1000111
|Copyright: © 2011 Suloma A, et al. This is an open-access article distributed under
the terms of the Creative Commons Attribution License, which permits unrestricted
use, distribution, and reproduction in any medium, provided the original author and
source are credited.
|The aim of the present study was to investigate the characteristics of gonadal fatty acid composition in 19 species
of wild coral reef fish (Serranidae, Lutjanidae, Lethridae, Siganidae and Labridae) from Philippine (11 species of
female) and Japanese (8 species of female and 5 species of male) waters with special attention to arachidonic acid
(ArA), docosahexaenoic acid (DHA) and eicosapentaenoic acid (EPA) levels and their ratios. ArA levels were always
higher than EPA levels in polar lipids of all the species and in neutral lipids in 17 of the 19 species. In ovarian polar
lipids of the 19 species, ArA level ranged from 6.0% to 19.4%, while EPA level ranged from 0.9% to 6.2%. Ovarian
DHA level was also always higher than EPA in all the species analyzed. Consequently, ArA/EPA ratios of these
species were high, unlike cold- and temperate-water species. ArA was the top fatty acid component in testis polar
lipids of three Lethrinus species (21.4% to 22.9%). Thus, ArA is not a minor component, that is, the major highly
unsaturated fatty acids (HUFAs) of polar lipids in all coral reef fish gonads are DHA and ArA (not EPA). The present
information on gonadal fatty acid composition can be used as a guideline for advancing appropriate broodstock diets
of tropical coral reef fish, emerging aquaculture commodities in developing countries.
|Arachidonic acid; EPA; DHA; Coral reef; Gonads; Fatty acids;
|Coral reef fishes are commercially very important. Some species
such as groupers and snappers are high-valued food fish and are
emerging as aquaculture commodities in tropical and subtropical
countries due to increasing demand . Fry availability is an essential
component in the development of the culture systems for new species
and in further increasing production of established culture species .
However, the expansion of aquaculture of coral reef fish is hindered by
unstable and limited supply of fry for aquaculture. At present, supply
of the fry is mostly dependent on natural resources, and the reduction
of the natural stock is a concern . In responding to this situation,
there has been considerable interest in developing reliable methods for
spawning and rearing larvae and fry. Spawning and egg/larval quality
are greatly affected by the quantity and quality of feed, and nutritional
deficiency could be one of the reasons for inconsistent quality of spawns
and fry. However, formulated broodstock diets for coral reef fish are still
under development because of the lack of information on nutritional
traits of these species.
|Marine fish have low or no capacity to synthesize highly
unsaturated fatty acids (HUFAs) from C18 fatty acids. HUFAs are
important components of cell membranes and are thought to play
important roles in membrane fluidity, modulation of enzyme activity,
neural development and regulation of stress resistance. Especially,
eicosapentaenoic acid (EPA: 20:5n-3) and docosahexaenoic acid
(DHA: 22:6n-3) are considered as dietary essential fatty acids for
normal growth and survival in most marine fish. Most of the studies
have focused on the effects of dietary supplementation of EPA and
DHA on broodstock performance. Indeed, dietary EPA and DHA
have successfully improved reproductive performance and egg/larvae
quality such as fecundity, embryo development, hatchability and
survival in several species [4,5]. Thus, the importance of EPA and DHA
in broodstock nutrition has been emphasized . Little attention was
given to the importance of dietary n-6 HUFA, especially arachidonic
acid (ArA: 20:4n-6) in marine fish, because ArA is found in only
small quantities in cold and temperate water fish, and dietary ArA was presumed to be unimportant in these species. Since Castell et al. 
found the dietary essentiality of ArA in juvenile turbot (Scophthalmus
maximus), the importance of ArA in fry production technologies of
marine fish has been re-considered .
|There have been some papers reporting that Australian or tropical
marine fish contain ArA levels equivalent to or higher than those of EPA
in muscle or edible parts [9-12] unlike cold and temperate water fish in
the Northern Hemisphere. However, this fact has been overlooked in
fry production of tropical or coral reef fish species. On the other hand,
Ogata et al. found that ArA is not a minor component in the ovaries
of several tropical species such as mangrove red snapper (Lutjanus
argentimaculatus), rabbitfish (Siganus guttatus and S. canaliculatus),
striped jack (Caranx fulvoguttatus) and coral trout (Plectropomus
leopardus), suggesting that this fatty acid may be nutritionally much
more important for egg development and larvae growth in the tropical
species than in cold and temperate water species . The degree of the
preferential retention of ArA in gonadal polar lipids appears larger in
tropical species than in cold and temperate water species. The degree
of the physiological importance of ArA in reproduction and larvae/fry
performance of tropical or coral reef fish might result in this difference
in HUFA profile between tropical and cold/temperate species. When
we consider the importance of dietary HUFA including ArA in fry
production, as described above, we should pay more attention to HUFA
characteristics in tropical fish.
|The aim of the present study was to investigate the characteristic of fatty acid composition of gonads in wild coral reef fish from Philippine
(11 species of female) and Japanese (8 species of female and 5 species
of male) waters, and special attention was paid to ArA, DHA and
EPA levels and their ratios. Gonadal fatty acid composition in fish is
affected by various biotic and abiotic factors, diets, species, size, age,
maturity, temperature, seasons, salinity etc. Nevertheless, HUFA profile
of gonads would be useful to develop broodstock diets for coral reef
species where their requirement data are not yet available. Although
the present information on gonadal fatty acid composition can be used
as a guideline for developing appropriate broodstocks diets for coral
reef fish, it should be noted that the conclusion of the present study
is based solely on limited data. Absolute comparison of HUFA profile
in coral reefs fish gonads would require much larger sample sizes and
more systematic sampling from different locations, seasons, species,
habitats and maturity.
|Materials and Methods
|Nineteen species of wild coral reef fish were obtained from local
dealers of three sites: Ishigaki (Okinawa, Japan), Puerto Princesa
(Palawan, Philippines) and Igang (Guimaras, Philippines). Fish were
immediately killed with iced water, transferred to local laboratories
with ice and directly dissected to take gonad samples. The sampled
gonads in Ishigaki were directly stored at -80°C for a night and sent to
our laboratory (Japan International Research Center for Agricultural
Sciences, Tsukuba, Japan) with dry ice by air, and thus fatty acid
composition in wet tissue was determined. Samples collected in
Philippines were temporarily stored at -20°C, being freeze-dried,
pulverized and sent by air. Fatty acid composition in dried tissue was
determined for the samples collected in Philippines. Species name,
body weight (g), body length (cm) and gonadosomatic index (GSI) were listed in Table 1. Of the samples, species names were not able to
be identified in two species of Epinephelus that were collected at Puerto
Princesa. The present paper gives them temporarily the names of
Epinephelus sp. 1 and Epinephelus sp. 2 (Table 1).
||Table 1: Fish name, body weight, body length and gonadosomatic index of sample fish (average±S.E.) and number of sample.
|All the wet and dried samples were stored at -80°C until lipid
extraction. Lipid extraction was carried out with a mixture of chloroform
and methanol (2:1, v/v)  containing 0.01% butylhydroxytoluene
(BHT). Total lipids were separated into polar (PL) and neutral lipids
(NL) with a silica cartridge (Sep-pak plus, Waters, Milford, MA, USA)
as described by  (Table 2). Fatty acid methyl esters (FAME) were
prepared by transesterification with borontrifluoride in methanol
according to the procedure of  and were purified with thin-layer
chromatography (Silicagel 70 Plate, Wako, Osaka, Japan; solvent system:
petroleum ether/diethyl ether/ acetic acid = 90:10:1, v/v). FAME were
analyzed using gas liquid chromatography (GC- 17A; Shimadzu, Kyoto,
Japan) equipped with a hydrogen flame ionization detector (FID) and
an Omegawax 320 fused silica capillary column (30 m x 0.32 mm i.d.;
Supelco, Bellefonte, PA, USA). The column temperature was initially
held at 160 C for 5 min, followed by an increase at a rate of 4 C/min
to a final temperature of 210°C. The carrier gas was helium and the
pressure was 80 kPa. Individual FAME was identified using a reference
standard (Funakoshi, Tokyo, Japan) and known fish meal FAME and
was quantified with an integrator (C-R7A plus; Shimadzu).
||Table 2: Composition of total lipid (%)*, neutral lipid (NL, % of total lipid) and polar lipid (PL, % of total lipid) (mean value±S.E.).
|Table 2 shows composition of total lipid (%, wet basis for Ishigaki
samples and dry basis for Philippine samples), neutral lipid (NL, % of
total lipid) and polar lipid (PL, % of total lipid). The data of Lethrinus
ornatus were not recorded due to a laboratory mistake. Total lipid levels ranged from 2.0% to 11.0% in ovaries (wet basis, Ishigaki), from 4.2%
to 28.0% in ovaries (dry basis, Philippines) and from 1.2% to 3.4% in
testes (wet basis, Ishigaki), respectively. The ranges of polar lipid levels
in ovaries and testes were respectively from 12.3% to 86.7% and from
43.7% to 74.8%, those of neutral lipid levels in ovaries and testes being
respectively from 13.3% to 87.7% and from 25.2% to 56.3%.
|Major fatty acids (% of total fatty acids) of the gonads in wild coral
reef fish are shown in Tables 3 for polar lipids and 4 for neutral lipids.
The most striking result in the present study was that ArA levels were
always higher than EPA levels in polar lipids of all the gonad samples
analyzed, and that in 17 of the 19 species, ArA levels were also higher
than EPA levels in neutral lipids of their gonads. In ovarian polar
lipids, ArA levels ranged from 6.0% (Epinephelus areolatus) to 19.4%
(Lethrinus atkinsoni), while EPA levels ranged from 0.9% (Siganus
virgatus) to 6.2% (Pristipomoide argyrogrammicus). The minimum
and the maximum ArA/EPA ratios in ovarian polar lipids were 1.3
in E. areolatus and 15.6 in Plectropomus leopardus (Japan). ArA in
ovarian polar lipids was one of the top three fatty acid components in
Cephalopholis argus (12.6% with an ArA/EPA ratio of 3.3), Epinephelus
quoyanus (15.2% with a ratio of 9.1), Epinephelus. sp.-1 (11.4% with
a ratio of 5.3), Epinephelus sp.-2 (11.7% with a ratio of 3.4), Lutjanus
decussatus (16.5% with a ratio of 5.3), Lutjanus erythropterus (17.5%
with a ratio of 4.1), P. leopardus (Japan) (19.2% with a ratio of 15.6),
Lutjanus gibbus (12.3% with a ratio of 4.3), Lethrinus miniatus (17.3%
with a ratio of 3.6), Siganus canaliculatus (Japan) (9.9% with a ratio of
4.7), L. atkinsoni (19.4% with a ratio of 4.1), Siganus guttatus (12.7%
with a ratio of 5.3) and S. virgatus (8.9% with a ratio of 10.0). Irrespective
of the maturity, thus, ovarian polar lipids had higher levels of ArA than
EPA levels, for instance in P. leopardus (Japan) (immature, Tables 1 and
3) and S. canaliculatus (mature, Tables 1 and 3). DHA was also one of
the top three fatty acid components in ovarian polar lipids in 15 of the
19 species. Ovarian DHA level in polar lipids was always higher than
EPA in all the species analyzed, ranging from 7.9% with DHA/EPA ratio
of 8.9 (S. virgatus) to 27.8% with a ratio of 18.3 (S. canaliculatus). DHA
levels in ovarian polar lipids were lower in the six species and higher in the remaining species than ArA levels, respectively. The variation of
DHA/ArA ratios with the range from 0.5 (P. leopardas, Japan) to 4.2
(E. areolatus) was relatively small among species, compared to those
of ArA/EPA (1.3-15.6) and DHA/EPA (3.0-18.3) ratios. Surprisingly,
ArA was the top fatty acid component in testis polar lipids of Lethrinus
ornatus (22.9%), Lethrinus nebulosus (22.5%) and L. atokinsoni (21.4%).
Consequently, ArA/EPA ratios of these species were extremely high
with the range from 8.4 to 20.2. ArA levels in testis polar lipids of two
Siganus species were intermediate but higher than EPA levels with the
ArA/EPA ratios of 10.0 and 6.1.
||Table 3: Major fatty acids (% of total fatty acids) in polar lipids of gonads of 19 species of coral reef fish from Philippines and Japanese water.
|ArA, EPA and DHA levels in neutral lipids (Table 4) were lower than
those in polar lipids due to relatively high levels of 16:0, 18:0 and 18:1n-
9 fatty acids in neutral lipids in gonad tissues. Yet, in neutral lipids, ArA
levels were entirely higher than EPA levels throughout all the species
with the exception of E. areolatus ovary and P. leopardus (Philippines)
ovary. ArA, EPA and DHA levels in neutral lipids of ovaries ranged
from 1.3% (Cephalopholis cyanostigma and E. areolatus) to 10.1% (S.
virgatus), from 0.5% (C. argus) to 4.9% (P. argyrogrammicus) and from
0.7% (C. argus) to 16.7% (P. argyrogrammicus), respectively. These
HUFA levels in testes ranged from 4.6% (L. ornatus) to 14.3% (L.
atokinsoni), from 1.3% (L. atokinsoni) to 2.8% (L. ornatus) and from
1.6% (L. ornatus) to 10.4% (S. guttatus), respectively.
||Table 4: Major fatty acids (% of total fatty acids) in neutral lipids of gonads of 19 species of coral reef fish from Philippines and Japanese water.
|It is obvious that ArA is not a minor HUFA but an essential
component in gonads of wild coral reef fish, judging from the results of
the present study and our previous paper that intermediate or high ArA
and DHA levels with relatively low EPA level were found in ovaries,
eggs, fry and/or muscle of mangrove red snapper (L. argentimaculatus),
two species of rabbitfish (S. guttatus and S. canaliculatus), striped jack
(C. fulvoguttatus) and coral trout (P. leopardus) sampled in Central
Philippines . Earlier papers reported that muscle tissue or edible
part of Australian [9,11], Malaysian  and Arabian Gulf  marine
fish were rich in ArA whose level was equivalent to or higher than
EPA level, unlike cold and temperate water species in the Northern Hemisphere. Mitochondrial membranes (liver and heart) of three coral
reef species had a significantly higher proportion of ArA than those
of cold water species, excluding Mugil cephalus (coastal and estuarine
species) . On the other hand, few papers have been available on
gonadal fatty acid composition of wild coral reef fish, especially in
commercially important species which are emerging as new aquaculture
commodities. Alava et al.  reported a relatively high ArA level in
wild-sourced grouper (Epinephelus coioides) broodstock (GSI: 0.73)
that the levels of HUFA were: DHA (13.1%) > ArA (5.1%) > EPA (1.9%)
with a DHA/EPA ratio of 6.8 and a DHA/ArA ratio of 2.5 in the ovarian
total lipids. Fatty acid compositions of ovaries, eggs and fry from not
wild but reared broodstock at tropical and subtropical hatcheries have been reported in striped mullet (Mugil cephalus) (coastal, estuarine and
bethopelagic species), milkfish (Chanos chanos) (coastal, estuarine
and bethopelagic species) , Asian seabass (Lates calcarifer) (coastal,
estuarine and demersal species) , white sea bream (Diplodus sargus)
(demersal and oceanodromous species)  and cobia (Rachycentron
canadum) (reef-associated pelagic species) . Since gonadal fatty
acid profile is greatly affected by feeds that have been fed during rearing
period, the results of the hatchery-raised broodstock are not directly
compared with the present ones based on wild coral reef fish. In general,
gonads and eggs of high and temperate latitude species in the northern
hemisphere have high levels of EPA and DHA, and consequently most of
these species show extremely low ArA/EPA ratios [24,25]. The average levels of ArA, EPA and DHA in egg polar lipids of seven Northwest
European species (wild) were 1.9%, 14.6% and 28.0%, respectively .
The average levels of ArA, EPA and DHA in ovarian polar lipids of
eight cold and temperate water species (wild) sampled in the Pacific
Ocean were 4.1%, 13.1% and 22.0%, respectively . In these species,
EPA levels were always higher than ArA levels with the exception of
Beryx splendens (bentopelagic species) in which ArA and EPA levels
were 5.4% and 4.6%, respectively (this specimen was sampled at 33°N
and 139°E). In the present result, however, ArA level was always higher
than EPA level in ovaries of all the species examined, irrespective of the
species, sampling location and a wide range of gonadosomatic index.
The average levels of ArA, EPA and DHA in ovaries of the coral reef
species were 12.3%, 3.4% and 17.2% in polar lipids and 3.9%, 1.8%
and 5.9% in neutral lipids, respectively. In this connection, eggs of six
Australian but freshwater or brackish water species (wild) have high or
middle DHA (8.2-29.3%) and ArA (1.8-15.3%) levels with low EPA
level (0.2-2.2%) in their polar lipids  as did the coral reef species.
We might say that coral reef fish appear to have a comparable ArA/EPA
ratio with freshwater fish rather than cold and temperate water marine
|Not only the concentration but also the mutual ratio of DHA, EPA
and ArA in diets may be important, and that the dietary optimum ratio
appears to vary depending on species, stage, the geography and the
species inhabits. In the present study, ovaries of 17 species of coral reef
fish had average ratios of: ArA/EPA of 4.8, DHA/EPA of 6.2 and DHA/
ArA of 1.7 in polar lipids and ArA/EPA of 2.5, DHA/EPA of 3.8 and
DHA/ArA of 2.5 in neutral lipids, respectively. This result indicates that
ovarian DHA/ArA ratio is about 2 in tropical coral reef fish, suggesting
that the DHA/ArA (not EPA) ratio of around or more than 2 may
be optimum for broodstock diets, especially for ovary development
in coral reef fish. We, therefore, recommend a dietary ratio of DHA/
ArA (not EPA) of about 2 or greater as an ideal for broodstock diets of
tropical coral reef fish.
|Information on testis fatty acid composition of wild fish has been
relatively scarce compared to ovaries and eggs. However, there have
been several papers reporting the effects of HUFA on male reproductive
performance in cold and temperate water species [27,28]. The present
study showed that ArA was the major HUFA in testis as well as in ovary
of the coral reef fish, especially in their polar lipids. Particularly, testis
ArA levels in polar lipids of the three Lethrinus species were more than
20%. The average levels of ArA, EPA and DHA in testis polar lipids of
nine species of cold and temperate water fish were 2.7%, 10.8% and
28.1%, respectively . The range of ArA, EPA and DHA levels in testis
polar lipids of three cold water marine species (Clupea pallasi, Theragra
chalcogramma and Osmerus eperlanus mordax) were from 1.2% to
4.6%, from 9.9% to 17.0% and from 14.1% to 20.9%, respectively .
In skipjack tuna Euthynnus pelamis (sampled in the Southern fishing
ground in the Pacific Ocean), testis polar lipids had ArA of 3.6%, EPA of
3.5% and DHA of 32.5%, respectively. It is also clear that the testes
of the coral reef fish analyzed here have higher ArA levels and ArA/
EPA ratios than those of cold and temperate water species. This result
suggests that ArA tends to be specifically much more concentrated in
testis of wild coral reef fish, and that the degree of the essentiality of
ArA during male reproductive process may be much greater in coral
reef fish compared to cold and temperate water species.
|Testes of the five species of coral reef fish had average ratios of: ArA/
EPA of 9.8, DHA/EPA of 6.5 and DHA/ArA of 0.8 in polar lipids and
ArA/EPA of 5.5, DHA/EPA of 3.4 and DHA/ArA of 0.6 in neutral lipids,
respectively. This result suggest that since the average DHA/ArA ratio was smaller in the testis than in the ovary, the optimum dietary ratio
of HUFA during gonadogenesis might be different between males and
females, and moreover that the degree of the physiological essentiality
of ArA might be greater in testis than in ovary.
|The high ArA levels in gonads observed in the present study shows
that coral reef fish species may require much more ArA for normal
gonadogenesis and embryo development than cold and temperate
water species, although dietary ArA tends to be concentrated in
gonads, eggs and sperm even in cold and temperate water species. ArA
as an eicosanoid precursor is well known to be multifunctional in fish
reproduction including pheromonal activity in mating behavior [31,32]: induction of ovulation , stimulation of steroidgenesis in ovary
 and testes , interaction of gonadotropin and prostaglandin
production . Coral reef fish have, in general, a longer spawning
period (from March through December) compared to cold and
temperate water species, and they spawn once a month during the
spawning period according to lunar phase. This mode of reproduction
might increase the degree of the essentiality of ArA in the reproduction
of coral reef fish compared with cold and temperate water fish. On
the other hand, tropical species hatch-out in a short time (usually less
than 24 h) during which the embryo has to build up many biological
systems, while cold water species such as salmonids take a longer time,
one or two months, to hatch out. Very fast embryo development in
tropical species might increase the degree of the essentiality of ArA,
prostaglandins and other eicosanoids. Heavy raining and typhoons
bring critical changes of temperature, salinity, irregular currents and
turbidity. The environmental stresses may further increase the degree
of the necessity for the eicosanoids, although the effectiveness of dietary
ArA appears different between the types of stress, acute (handling
stress) and chronic (salinity change) .
|It is widely accepted that fatty acid composition of fish tissues
reflects dietary fatty acid composition. It is also obvious that ArA level
in tissues including gonads is affected by dietary ArA level. When we
consider the fact that most of marine fish lack the de novo ability to
produce HUFA, the coral reef fish should also rely on a dietary supply
of HUFA from the food web. EPA and DHA in fish have been known
to be derived from the pelagic food chain from phytoplankton to
accumulate in higher order carnivores, especially in pelagic species and
deeper offshore demersal species . All the species investigated in
the present study were reef-associated species. Serranidae, Lujanidae,
Lethridae and Labridae are carnivores, which feed on smaller fishes,
crabs, shrimps, cephalopods, polychaeta worms, gastropods and
urochordates. Sigaidae are herbivores, which feed mainly on benthic
macroalgae. The coral reef fish perhaps depend heavily on their natural
food as their ArA sources (without doubt), although there might be a
special mechanism by which they store much more preferentially ArA
in their gonads than cold and temperate water fish.
|Renaud et al.  investigated fatty acid composition of 18 species
of tropical Australian microalgae . Only two pelagic species,
Nitzschia sp. and Fragilarias sp. had relatively high ArA levels equivalent
to EPA levels, but in the remaining 16 species, EPA was the major
HUFA and ArA level was low. EPA was also the major HUFA in tropical
phytoplankton sampled from coastal waters of the South China Sea
during one year cycle . In tropical paracalanid copepods, EPA level
is higher than ArA level with DHA/EPA/ArA ratios of 14:3:1 for Acartia
sinjiensis, 20:9:1 for Parvocalanus crassirostris and 25:6:1 for Bestiolina
similes, respectively . Thus, planktonic microorganisms do not
appear to be the primary source of ArA even in tropical waters. High
ArA levels are found in some species of marine red and brown seaweeds from both temperate and tropical waters, although this phenomenon is
not always limited to tropical areas [42-44]. Nevertheless, red and brown
seaweeds may be at least one of the sources. Dunstan et al. found
that in temperate marine fish from Southern Australian coastal waters,
demersal omnivore species (macroalgae consumers) have relatively high
ArA/EPA ratio (0.9) compared to demersal carnivores (0.6) and pelagic
carnivores (0.2) . Thus, ArA appears to be provided primarily from
some organisms existing in/on benthic substrate and benthic detritus
rather than pelagic organisms. This speculation may be supported by a
finding by Svetashev et al.  that ArA content is remarkably higher
in tropical holothurians than in the temperate species . Scarce
data are available on fatty acid composition of benthic prokaryotes
and eukaryotes, bacteria, fungi and protozoa towards the beginning of
ArA source in tropical marine food chain. The present result, high ArA
levels in coral reef fish, suggests that the existence of an ArA-rich food
chain may be widespread in coral reef areas, and that the widespread
existence of ArA-rich food chain may lead to comparatively higher ArA
contents in the coral reef fish. However, the issue of ArA origin in the
coral-reef food web is still unclear.
|Effort is needed to establish formulated diets appropriate for
broodstock of coral reef fish. Feeding locally available trash fish rich in
ArA would be at least one of the measures for improving broodstock
performance in coral reef fish, until appropriate formulated diets are
established for coral reef broodstock. Nevertheless, the information in
the present study can be used as a guideline (a dietary ratio of DHA/
ArA of about 2 or greater) for development of appropriate broodstock
and larval diets, to ensure high egg and larval quality of sustainable
hatchery production in tropical and subtropical areas. We have already
initiated a follow-up study on the effects of dietary ArA on reproductive
performance and larval/fry quality in coral reef fish. Results of feeding
trials will be published in separate papers.
|We would like to thank Dr. K.D. Shearer, Northwest Fisheries Science Center,
NMFS, Seattle, WA USA, for his critical reading of the manuscript. The authors are
also grateful to Mr. D.R. Chavez, INVE Asia Services Ltd, Thailand, Mr. E.S. Garibay,
the Aquaculture Department, SEAFDEC, Philippines and Dr. H. Furuita, National
Research Institute of Aquaculture, FRA, Japan for their technical assistants for
conducting the present study. This study was supported by the collaborative project
titled "Studies on Sustainable Production Systems of Aquatic Animals in Brackish
Mangrove Areas" between Japan International Research Center for Agricultural
Sciences, Japan and the Aquaculture Department, SEAFDEC, Philippines.
- Johnston B (2007) Economics and marked analysis of the live reef-fish trade
in the Asia-Pacific region. ACIAR Working Paper No.63, Australian Centre for
International Agricultural Research, Canbera.
- Marte CL (2003) Larviculture of marine species in Southeast Asia: current
research and industry prospects. Aquaculture 227: 293-304.
- Tupper M, Sheriff N (2008) Capture-based aquaculture of groupers. In: A.
Lovatelli, P.F. Holthus (eds), Capture-based aquaculture: Global overview, FAO
Fisheries Technical Paper 508, Food and Agriculture Organization of the United
Nations, Rome, pp. 217-253.
- Watanabe T, Arakawa T, Kitajima C, Fujita S (1984) Effect of nutritional quality
of broodstock diets on reproduction of red sea bream. Nippon Suian Gakkaishi
- Izquierdo MS, Fernández-Palacios H, Tacon AGJ (2001) Effect of broodstock
nutrition on reproductive performance of fish. Aquaculture 197: 25-42.
- Sargent JR (1995) Origin and functions of eggs lipids: nutritional implications.
In: N.R. Bromage, R.J. Roberts (eds), Broodstock Management and Egg and
Larval Quality, Blackwell Science, London, pp. 353-372.
- Castell JD, Bell JG, Tocher DR, Sargent JR (1994) Effects of purified diets
containing different combinations of arachidonic and docosahexaenoic acid on
survival, growth and fatty acid composition of juvenile turbot (Scophthalmus
maximus). Aquaculture 128: 315- 333.
- Bell JG, Sargent JR (2003) Arachidonic acid in aquaculture feeds: current
status and future opportunities. Aquaculture 218:491-499.
- Gibson RA (1983) Australian fish - an excellent source of both arachidonic acid
and ω3 polyunsaturated fatty acids. Lipids 18: 743-752.
- Gibson RA, Kneebone R, Kneebone GM (1984) Comparative levels of
arachidonic acid and eicosapentaenoic acid in Malaysian fish. Comp Biochem
Physiol C 78: 325-328.
- Fogerty AC, Evans AJ, Ford GL, Kennett BH (1986) Distribution of ω6 and ω3
fatty acids in lipid classes in Australian fish. Nutrition Reports International 33:
- Rawdah TN, El-Faer MZ (1994) Fatty acid composition of three commercially
important fish of the Arabian Gulf. Food Chem 51:193-196.
- Ogata HY, Emata AC, Garibay ES, Furuita H (2004) Fatty acid composition
of five candidate aquaculture species in Central Philippines. Aquaculture 236:
- Folch J, Lees M, Sloane-Stanley GH (1957) A simple method for the isolation
and purification of total lipids from animal tissues. J Biol Chem 226: 497-509.
- Juaneda P, Rocquelin G (1985) Rapid and covenient separation of
phospholipids and non phosphorous lipids from rat heart using silica cartridges. Lipids 20: 40-41.
- Miyashita K, Inukai N, Ota T, Sasaki S, Ota T (1999) Antioxidant activity of
water extracts from fish eggs on PC liposomes. Nippon Suisan Gakkaishi 65:
- Irving DO, Watson K (1976) Mitochodrial enzymes of tropical fish: A comparison
with fish from cold-waters. Comp Biochem Physiol B 54: 81-92.
- Alava VR, Priolo FMP, Arnaiz M, Toledo JD (2004) Amino and fatty acid profiles
of wild-sourced grouper (Epinephelus coioides) broodstock and larvae. In: M.A.
Rimmer, S. McBride, K.C. Williamas (eds) , Advances in Grouper Aquaculture,
ACIAR Monograph 110, Canbera, pp. 53-54.
- Tamaru CS, Ako H, Lee C-S (1992) Fatty acid and amino acid profiles of
spawned eggs of striped mullet, Mugil cephalus L. Aquaculture 105: 83-94.
- Ako H, Tamaru CS, Lee C-S (1994) Chemical and physical differences in
milkfish (Chanos chanos) eggs from natural and hormonally induced spawns. Aquaculture 127: 157-167.
- Nocillado JN, Peñaflorida VD, Borlongan, IG (2000) Measures of egg quality
in induced spawns of the Asian sea bass, Lates calcarifer Bloch. Fish Physiol
Biochem 22; 1-9.
- Cejas JR, Almansa E, Villamandos JE, Badía P, Bolaños A, et al. (2003) Lipid
and fatty acid composition of ovaries from wild fish and ovaries and eggs from
captive fish of white sea bream (Diplodus sargus). Aquaculture 216: 299-313.
- Faulk C, Holt GJ (2005) Advances inrearing cobia Rachycentron canadum larvae in recirculating aquaculture systems: Live prey enrichment and
greenwater culture. Aquaculture 249: 231-243.
- Tocher DR, Sargent JR (1984) Analyses of lipids and fatty acids in ripe roes of
some Northwest European Marine Fish. Lipids 19: 492-499.
- Takama K, Suzuki T, Yoshida K, Arai H, Anma H (1994) Lipid content and fatty
acid composition of phospholipids in white-flesh species. Fisheries Science 60:
- Anderson AJ, Arthington AH, Anderson S (1990) Lipid classes and fatty acid
composition of the eggs of some Australian fish. Comp Biochem Physiol B 96:
- Bell MV, Dick JR, Thrush M, Navarro JC (1996) Decreased 20:4n-6/20:5n-3
ratio in sperm from cultured sea bass, Dicentrarchus labrax, broodstock
compard with wild fish. Aquaculture 144: 189-199.
- Asturiano JF, Sorbera LA, Carrillo M, Zanuy S, Ramos J,et al,(2001) Reproductive performance in male European sea bass (Dicentrarchus labrax,
L.) fed two PUFA-enriched experimental diets: a comparison with males fed a
wet diet. Aquaculture 194: 173-190.
- Ota T, Takagi T (1989) Fatty acids of lipids from fish testes with particular
reference to furan fatty acids. Bulletin of the Faculty of Fisheries Hokkaido
University 40: 193-201.
- Hiratsuka T, Kitagawa T, Matsue Y, Hashidume M, Wada S (2004) Lipid class
and fatty acid composition of phospholipids from the gonads of skipjack tuna . Fisheries Science 70: 903-909.
- Sorensen PW, Goetz FW (1993) Pheromonal and reproductive function of
F-prostaglandins and their metabolites in teleost fish. J Lipid Mediat 6: 385-393.
- Ogata H, Kitamutra S, Takashima F (1994) Release of 13, 14-dihydro-ketoprostaglandin
F2a, a sex pheromone, to water by cobitid loach following
ovulatory stimulation. Fisheries Science 60: 143-148.
- Stacey NE, Goetz FW (1982) Role of prostaglandins in fish reproduction. Canadian Journal of Fisheries and Aquatic Sciences 39: 92-98.
- van der Kraak G, Chang JP (1990) Arachidonic acid stimulates steroidgenesis
in goldfish preovulatory ovarian follicles. Gen Comp Endocrinol 77: 221-228.
- Wade MG, van der Kraak G (1993) Arachidonic acid and prostaglandin
E2 stimulate testosterone production by goldfish testis in vitro. Gen Comp
Endocrinol 90: 109-118.
- Asturiano JF, Sorbera LA, Zanuy S, Carriolla M (2000) Effects of polyunsaturated
fatty acids and gonadtropin on prostaglandin series E production in a primary
testis cell culture system for the European sea bass. J Fish Biol 57: 1563-1574.
- Koven W, van Anholt R, Lutzky S, Atia IB, Nixon O, et al. (2003) The effect of
dietary arachidonic acid on growth, survival, and cortisol levels in different-age
gilthead seabream larvae (Sparus auratus) exposed to handling or daily salinity change. Aquaculture 228: 307-320.
- Sargent JR, Whittle KJ (1981) Lipids and hydrocarbons in the marine food web. In: A.R. Longhurst (eds), Analysis of Marine Ecosystems, Academic Press,
London, pp. 491-533.
- Renaud SM, Thinh L-V, Parry DL (1999) The gross chemical composition
and fatty acid composition of 18 species of tropical Australian microalgae for
possible use in mariculture. Aquaculture 170: 147-159.
- Shamsudin L (1998) Seasonal variation of fatty acid content in natural
microplankton from the Tumpat coastal waters of the South China Sea. Arch
Physiol Biochem 106: 253-260.
- McKinnon AD, Duggan S, Nichols PD, Rimmer MA, Semmens G, et al. (2003)
The potential of tropical paracalanid copepods as live feeds in aquaculture. Aquaculture 223: 86-106.
- Johns RB, Nicholas PD, Perry GJ (1979) Fatty acid composition of ten marine
algae from Australian waters. Phytochemistry 18: 799-802.
- Dembitsky VM, Pechenkina-Shubina EE, Rozentsvet OA (1991) Glycolipids
and fatty acids of some seaweeds and marine grasses from the Black Sea. Phytochemistry 30: 2279-2283.
- Vaskovsky VE, Khotimchenko SV, Xia B, Hefang L (1996) Polar lipids and fatty
acids of some marine macrophytes from the Yellow Sea. Phytochemistry 42:
- Dunstan GA, Sinclair AJ, O'Dea K, Naughton JM (1988) The lipid content and
fatty acid composition of various marine species from Southern Australian
coastal waters. Comp Biochem Physiol B 91: 165-169.
- Svetashev VI, Levin VS, Lam CN, Nga DT (1991) Lipid and fatty acid composition
of holothurians from tropical and temperate waters. Comp Biochem Physiol B