You are on page 1of 22

COMPARISON OF FATTY ACIDS (FAS)

PROFILE OF HELICOLENUS
DACTYLOPTERUS, SYNAGROPS MICROLEPIS
& CHLOROPHTHALMUS AGASSIZI, OFF
NAMIBIA.

KANYIKI VILHO ROYAL


2017

BACHELOR OF SCIENCE (HONOURS) IN FISHERIES AND AQUATIC SCIENCES


THE UNIVERSITY OF NAMIBIA

SUPERVISOR: DR. IITEMBU JOHANNES


Abstract
The Benguela ecosystem is important to the fishery off Namibia. Helicolenus dactylopterus,
Synagrops microlepis & Chlorophthalmus agassizi are important species in the Benguela
ecosystem. The three species are found in the diet of many predatory fish species in marine
waters off Namibia, live at an overlapping depth ranges and have common prey in their diets.
The objective of this study was to compare the fatty acid profiles of these three species.
Multivariate tests revealed significant differences in the storage of MUFA, PUFA and SFA
profile between the three species, displaying that fatty acids in fish differ between species. All
the species had higher PUFA compared to MUFA and SFA, agreeing that Polyunsaturated
Fatty Acids are the most active and dominant fatty acids in marine fish. This indicate that
although these species have prey in their difference, there significant differences is in their
dietary source.
Keywords: Fatty Acids, MUFA, PUFA, SFA, Helicolenus dactylopterus, Synagrops
microlepis & Chlorophthalmus agassizi.

[i]
Table of contents
Abstract .................................................................................................................................................. i
List of Tables ....................................................................................................................................... iii
List of Figures ...................................................................................................................................... iii
List of Abbreviation ........................................................................................................................... iii
1. Introduction ..................................................................................................................................... 1
2. Material and Methods .................................................................................................................... 3
2.1 Study area and field sampling .................................................................................................... 3
2.2 Laboratory analysis..................................................................................................................... 4
2.3 Statistical analyses ...................................................................................................................... 4
3. Results ............................................................................................................................................... 5
3.1 FA profile...................................................................................................................................... 5
3.2 MonoUnsaturated Fatty Acids.................................................................................................... 7
3.3 PolyUnsaturated Fatty Acids ...................................................................................................... 7
3.4 Saturated Fatty Acids .................................................................................................................. 7
4. Discussion......................................................................................................................................... 8
5. Reference ........................................................................................................................................ 11
6. Appendices ..................................................................................................................................... 15

[ii]
List of Tables
Table 1. The final 30 FAs detect in amount above 1%.........................................................5

List of Figures
Figure 1. The sampling stations……………………………………………………………3

Figure 2. Mean value of MUFA, PUFA and SFA…………………………………............6

List of Abbreviation

FA Fatty acids
PUFA Polyunsaturated fatty acids
MUFA Monounsaturated fatty acids
SFA Saturated fatty acids
DHA Docosahexaenoic acid
EPA Eicosapentaenoic acid
MANOVA Multivariate analysis.

[iii]
1. Introduction
The Benguela ecosystem which extend over three countries (South Africa, Namibia and
Angola) is one of the coastal upwelling systems in the world (Coetzee et al. 2008). High
phytoplankton productivity in the nutrient rich upwelling waters is the basis for the highly
productive Benguela ecosystem. The Benguela ecosystem is important to the fishery
communities off Namibia, it supports wide range of marine fishery with catches and production
of commercial species of over a million tons per year, (Sakko 1998). Studies conducted in the
Benguela ecosystem allow better understanding of liveliness of marine species and contribute
knowledge toward their diverse exploration, hence find strategic managements that will sustain
and protect their population. Jacopever (Helicolenus dactylopterus), Thinlip splitfin
(Synagrops microlepis) and Shortnose greeneye (Chlorophthalmus agassizi) fish are important
species in the Benguela ecosystem. They are studied to live at an overlapping depth ranging
from 100 to 1000m deep, (Anastasopoulou et al. 2005; Hamukuaya et al. 2001; Sequeira et al.
2009). The Jacopever is a benthic deep-water fish which is widespread in the eastern and north
Atlantic Ocean, (Sequeira et al. 2009). Shortnose greeneye is a demersal species that live in
mud and clay bottoms and is very abundant in the Western Atlantic Ocean, (Anastasopoulou
et al. 2005). The Thinlip splitfin is a bathypelagic species normally found in the Eastern
Atlantic off the coasts of Walvis Bay, (Hamukuaya et al. 2001). These three Benguela species
have common prey in their diets, both benthic and pelagic organisms such as crustaceans,
fishes, cephalopods, and echinoderms, (Eschmeyer and Dempster 1990). Chlorophthalmus
agassizi however showed a mixed feeding strategy, exploiting a wide range of prey including
mesopelagic, benthic and endo-benthic organisms, (Anastasopoulou and Kapiris 2007). The
three species are found in the diet of many predatory fish species (e.g. Hake species) in marine
waters off Namibia. Literatures indicates early exploration of these species, for example
Macpherson & Roel (1987) studied the daily ration and feeding periodicity of some fishes off
the coast of Namibia and trophic relationships in the demersal fish community off Namibia.
The findings of the above cited studies were based on stomach-content analyses. Therefore, the
use of Fatty acids analysis have improved our understanding of many marine ecological
relationships, they provide long-term and time-integrated dietary information about consumers,
(Koussoroplis et al. 2010). Fatty acids have previously been used to examine qualitative aspects
of food webs, energy transfers, and predator prey relationships, (Sara et al. 2004). Hence the
method of exploring consumer diets based on fatty acid profiles represents an additional and
complementary approach to those already used and may shed further light on the trophic

[1]
dynamics of closely related species that inhabit the same oceanic regions, (Iitembu and
Richoux 2014). The difference of fat acids compositions of marine fish depends on diet,
geographic conditions, body length, sex, species, and fat content, (Baris et al. 2014).
Remarkably, fatty acids have been used as qualitative markers to trace or confirm predator-
prey relationships in the marine environment for more than thirty years, (Dalsgaard et al. 2003).
Fatty acids are also known to play a number of key roles in metabolism (storage and transport
of energy), as essential components of all membranes, and as gene regulators (Drevon 2010).
As part of complex lipids, fatty acids can be saturated, monounsaturated or polyunsaturated
(Drevon 2010). Robin et al. (2003) studied that the fatty acid (FA) content of fish reflect fatty
acid composition of the diet. Hence the incorporation of FA into tissues is modulated by various
metabolic factors, and final composition will depend upon the initial FA content, cumulative
intake of dietary fatty acids, growth rate and duration, (Robin et al. 2003). The objective of this
study was to compare the fatty acid profiles of these three species. The specific objectives of
the study were to compare fatty acid profiles within and between the species. This study
therefore contributes towards research effort of understanding their interspecific trophic
relationships. Knowledge from this study is useful toward sustainable fisheries practice
especially in understanding prey-predatory species interactions.

[2]
2. Material and Methods

2.1 Study area and field sampling

Data used in this study were collected by the Ministry of Fisheries and Marine resources, during
demersal survey (2012). Sample were collected from pre-determined survey stations. The
collections were completed during a hake biomass survey (11 January–25 February 2011) on
board MV Blue Sea I and a monkfish biomass survey (16–27 December 2011) on board RV
Welwitschia. Sampling for all fish was opportunistic, with the general aim of obtaining a wide
size distribution of each species. At each station, 1–10 individuals (depending on availability)
were selected, (see Iitembu and Richoux 2016).

Figure 1. Sampling stations along the Atlantic Namibian coastline

[3]
2.2 Laboratory analysis

The laboratory analyses of the FA extract were done at Rhodes University. A small section of
white muscle was removed from the anterodorsal region of each frozen fish, then lyophilized
at −60 °C for 24 hour. The Lyophilized samples were first grounded individually with a mortar
and pestle into a fine powder then placed in a glass test tube, after which chloroform (with
butylated hydroxytoluene) was added, the tube were later flushed with nitrogen and then stored
at −20 °C. Total lipid were extracted in 8:4:3 (v/v/v) of CHCl3/methanol/water. A neutral lipids
of tissues were extracted from the total lipids using column chromatography on silica gel, (See
Iitembu and Richoux 2016).

2.3 Statistical analyses

MANOVA was used to determine whether there is significant differences in the Fatty acids
profiles of the three species. The Tukey HSD was used to determine the Fatty acids that
contributed more to the significant difference of the FA profiles between the three species.

[4]
3. Results

3.1 FA profile

A total of 24 samples of the three species collected along different station off the Benguela
region were analyzed for fatty acid (FA) profiles. Fifty three (53) FAs ranging from 14 to 24
carbon atoms in length, were identified from the samples. Analyses was limited to most
abundant 30 FAs (Table 1) that were detected in amounts >1 % in all species. All the species
had higher PolyUnsaturated Fatty Acids (PUFA) compared to MonoUnsaturated Fatty Acids
(MUFA) and Saturated Fatty Acids (SFA) (Fig 2). H. dactylopterus showed higher PUFA and
SFA content compared to other species while C. agassizi had a higher MUFA content (Fig 2).
A significant difference between the MUFA, PUFA and SFA profiles of the three species was
observed, (p<0.05), (p<0.05) and (p<0.05).
There was a significant differences between the FA profiles of the three species (MANOVA,
Wilk’s λ=33.22, p<0.05). A Tukey HSD of the FAs indicated that 14:0 was significantly
different between H. dactylopterus and S. microlepsis (p<0.05) and between C. agassizi and
H. dactylopterus (p<0.05). 18:1w7 was significantly different between C. agassizi and H.
dactylopterus (p<0.05) and between H. dactylopterus and S. microlepsis (p<0.05). Tukey HSD
indicated a significant difference in 22:6w3 of C. agassizi and H. dactylopterus (p<0.05), and
between H. dactylopterus and S. microlepsis (p<0.05).

Table 1. The 30 FAs detect in amount above 1% used in the parametric analysis.

FAs Chlorophthalmus agassizi Helicolenus dactylopterus Synagrops microlepsis


(n=12). 12-20 cm (n=9). 12-14 cm (n=3). 13-15 cm
Mean(SD) Mean(SD) Mean(SD)
16:1w7 .48(1.17) .33(.65) 1.79(2.71)
17:1w7 .38(.26) .39(.22) .38(0.87)
18:1w9 15.07(10.37) 7.58(6.23) 25.91(15.07)
18:1w7 11.19(14.50) .79(1.44) 3.19(.24)
18:1w5 .69(1.23) .059(.11) 13.68(23.41)
20:1w9 3.27(2.93) 1.92(1.92) 4.70(4.14)
22:1w9 3.64(2.24) .64(.89) 3.26(.37)

[5]
20:4w3 .55(.23) 4.09(1.52) 14.03(.06)
20:5w3 9.01(2.51) .07(.13) .56(.06)
20:2w6 .20(.10) .09(.19) .18(.02)
20:3w6 .03(.06) .08(.23) 13.01(.02)
20:4w6 1.73(1.08) 4.10(1.53) .63(.08)
20:3w3 .08(.08) .07(.13) 9.04(.06)
21:5w3 .00(.00) .17(.41) 5.53(.59)
18:2w6 .18(.36) .32(.50) .00(.00)
18:4w3 .40(.42) .00(.00) .00(.00)
22:2 .00(.00) 8.24(2.75) .00(.00)
22:4w6 .17(.12) .48(.47) 8.04(.06)
22:5w6 .44(.29) 1.24(.55) .17(.01)
22:5w3 2.35(.94) 3.29(1.43) 1.62(.17)
22:6w3 24.43(14.31) 39.10(7.36) 8.24(.99)
14:0 5.13(2.44) 1.25(.65) 7.83(.82)
15:0 .314(.12) .43(.18) .28(.02)
16:0 15.29(11.49) 21.73(2.34) 15.59(13.53)
(i)17:0 .29(.15) .41(.34) .30(.05)
17:0 .62(.17) .63(.27) .48(.05)
18:0 3.22(2.47) 5.90(2.25) 4.14(3.59)
20:0 .25(.12) .29(.65) .37(.04)
(i)15:0 .12(.07) .07(.06) .13(.01)
(ai)7:0 .09(.06) .16(.26) .09(.01)
21:0 .08(.11) .02(.06) .08(.09)

700
Average FAs composition (%)

600

500

400

300

200

100

0
MUFA PUFA SFA

Chlorophthamus agassizi Helicolenus dactylopterus Synagrops microlepsis

Figure 2. Average composition of MUFA, PUFA AND SFA, observed from the 30 FAs.

[6]
3.2 MonoUnsaturated Fatty Acids

There was significant difference between the MUFA profiles of the three species (MANOVA,
Wilk’s λ= 0.129, p<0.05). A Tukey HSD indicated that 18:1w9 was significantly different
between H. dactylopterus and S. microlepsis, (p<0.05); 18:1w7 was significantly different
between C. agassizi and H. dactylopterus (p<0.05) and 22:1w9 was significantly different
between H. dactylopterus and C. agassizi, (p<0.05) and between H. dactylopterus and S.
microlepsis (p<0.05).

3.3 PolyUnsaturated Fatty Acids

There was significant difference between the PUFA profiles of the three species (MANOVA,
Wilk’s λ= 0.044, p<0.05). A Tukey HSD indicated that 20:4w3, 20:5w6 and 22:2 were
significant different between C. agassizi and H. dactylopterus, (p<0.05), (p<0.05) and
(p<0.05); 22:6w3 was significantly different between S. microlepsis and H. dactylopterus,
(p<0.05) while Fatty acid 22:5w3 was significantly different between C. agassizi and S.
microlepsis, (p<0.05).

3.4 Saturated Fatty Acids

There was significant difference between SFA profiles of the three species (MANOVA, Wilk’s
λ= 0.101, p<0.05). A Tukey HSD indicated that Fatty acid 14:0 was significantly different
between the SFAs of C. agassizi and H. dactylopterus, (p<0.05) and between S. microlepsis
and H. dactylopterus (p<0.05) while 15:0 was significantly different between S. microlepsis
and C. agassizi (p<0.05).

[7]
4. Discussion

This study aimed at comparing the fatty acid profiles of Helicolenus dactylopterus, Synagrops
microlepsis and Chlorophthamus agassizi. Multivariate tests revealed significant differences
in the storage of MUFA, PUFA and SFA profile between the three species. All the species had
higher PUFA compared to MUFA and SFA. H. dactylopterus showed higher PUFA and SFA
content compared to other species while C. agassizi had a higher MUFA content.

A mean comparison between the species FAs indicated a higher PUFA than MUFA and SFA
composition. The composition of the fatty acids in fish differ between species (Jalaludin 2013),
depending on structure, properties, requirements and functions in the body (Baeza 2015). The
results had reflected similarly with Baeza (2015) findings, the aquatic medium is characterized
by a wealth of PUFAs with fish containing always an elevated percentage of PUFAs. The
higher storage of PUFA in fish is general reflected by its demonstrated major function in
different aspects of the fish development, (Field 2003). They have been shown to alter the
expression of numerous genes involved in the metabolic function of the cell, modulate the
expression of a variety of genes coding for key regulatory proteins in metabolic pathways such
as those involved in digestion, glycolysis, glucose transport, inflammation, and cellular
communications, (Field 2003).
Palou (2007) highpoint Polyunsaturated Fatty Acids (PUFA) as most active and dominant fatty
acids in marine fish. This is acknowledged for their role as physiologically active factor in
many fish species to actively participate in gonad maturation, egg quality (Izquierdo et al.
2001) and larval growth of fish (Tulli and Tibaldi 1997).
Comparably, SFAs and MUFAs are heavily catabolized for energy in fish because they are
consumed in large amounts during growth, (Baeza 2015). They are identified in the body of
fish to be more structural in nature, and thus more rapidly influenced by changes in the
requirement of metabolic energy, (Koop- man et al. 1996, Iverson et al. 2002). Therefore, the
mobilization of SFA is required solely for provision of metabolic energy and less destined in
gonads and larval development of the fish, (Sargent et al. 1989). While MUFAs are crucial
stored to provide special conformational properties to the bio-membranes, and assist tissue
specific cells in reacting to external stimuli such as changing environmental temperatures and
light regimes (Sargent et al. 1993, Cook 1996).

[8]
The results agrees with other studies on FAs, most marine fish lack delta -5 desaturase activities
needed to biosynthesize PUFAs and therefore have an absolute dietary requirement for
unsaturated FAs (Tocher and Ghioni 1999; Hastings et al. 2001; Nichols 2003). Marine species
present low enzymatic activity and depend almost completely on their diet to obtain the main
long-chain n-3 PUFAs, (Toucher 2010). Hence, unsaturated fatty acids may be synthesized by
animals but only to a limited extent and must be largely supplemented by the diet (Steffens,
1997).
H. dactylopterus showed higher PUFA and SFA contents compared to other species. Although
very little information is available on the feeding habits of this species, studies indicated that
the species feeding strategy which, according to Macpherson (1985), is primary a daytime
predator feeding during a relative short period. Also a two clear dietary shifts (at 20 cm and 28
cm TL) occur along H. dactylopterus life, (Sequeira et al. 2009). Nutritional needs during the
species growth changes, large individuals show major consumption of natantia which are rich
source of SFA, (Sequeira et al. 2009).

C. agassizi had a higher MUFA content compared to other species. The species is studied to
be an active carnivorous predator of benthic, an agile prey that exhibits a high feeding activity
(Anastasopoulou & Kapiris 2007). Stomach content analysis indicated that fish and crustacean
are abundant prey but a high abundance of plant detritus (plant sources of fat tend to be very
high in monounsaturated and polyunsaturated fats, (Assy et al. 2010)) found in their stomachs,
confirmed a mixed type of diet, (Anastasopoulou & Kapiris 2007).

A significant differences was found between the MUFAs of the three species. Asclepic acid
(18:1w7) and Eurisic acid (22:1w9) were significantly different between H. dactylopterus and
C. agassizi. Asclepic and Eurisic acids belongs to the class of chemical entities known as long-
chain fatty acids, these are fatty acids with an aliphatic tail that contains between 13 and 21
carbon atoms (Lambertsen 1977). Studies (Assy et al. 2010) outlined that some marine
organisms lack the ability to introduce double bonds in long-chain fatty acids beyond carbon 9
and 10, hence this might had considerable influenced the significant difference between the
different species.

There was significant difference between the PUFA profiles of the three species. DHA
(22:6w3) was significantly different between S. microlepsis and H. dactylopterus. Fatty acid
such as EPA and DHA that been found in marine fish are originally obtain from the
phytoplankton and also seaweed that include in their food chain (Cavington 2004). Therefore,

[9]
the difference in DHA could be a result of species different feeding activities as studies
indicates that the S. microlepsis migrate towards the surface at night for feeding purpose
(Heemstra 1984) which allow it to feed on the numerous floating populations of plankton
species which contain significant concentrations of EPA and DHA.

A significant difference was found between the SFA profiles of the three species. Myristic acid
(14:0) was significantly different between H. dactylopterus and S. microlepsis. Myristic acids
are found in all fish fats, and probably originate in marine phytoplanktonic algae (Lambertsen
1977). S. microlepsis storage of relatively high fat content (Iitembu and Richoux 2014) could
mean a high incorporation of 14:0 SFA hence reflect the difference in the storage between the
different species.
The findings of the study has shed light on the exploration of fish feeding based on fatty acids
(FAs). As observed, the species has more influence on the difference of fat acids compositions
between marine fish with additions to diet, geographic conditions, body length, sex and fat
contents. Studies have established that specific habits and characteristics from different fish
families, such as nocturnal/diurnal habits, anatomical and physiological characteristics
generally present differences in intestinal enzyme secretion metabolism, which influence
nutrient digestion and absorption processes, (Logato 1998). Thus, in addition to fish characters,
genetic potential and fatty acid enzymatic biosynthesis, can influence the final FA profile in
muscle tissue which can vary between species, (Toucher 2003).

More advanced and directed scientific studies must be conducted on the Benguela ecosystem
to document and cover all the important aspects of the environment interactions from fish to
planktons. Fish exploration is key to sustainable exploitation and truthful management
practices of marine environments. Hence, further studies should be developed were Stomach
contents analysis can be used together with fatty acids as quantitative and qualitative markers
to trace or confirm predator-prey relationships in the marine environment.

[10]
5. Reference

Anastasopoulou A. & Kapiris K. 2007. Feeding ecology of the Shortnose greeneye


Chlorophthalmus agassizi (Bonaparte 1840) in the eastern Ionian Sea (eastern
Mediterranean). Journal of Applied Ichthyology 24: 170–179.

Anastasopoulou A, Yiannopoulos C, Megalofonou P & Papaconstantinou C. 2005. Distribution


and population structure of the Chlorophthalmus agassizi (Bonaparte, 1840) on an
unexploited fishing ground in the Greek Ionian Sea. Journal of Applied Ichthyology 22:
521–529.

Assy N, Grosovski M. 2010. Monounsaturated Fat Enriched with Olive Oil in Non-alcoholic
Fatty Liver Disease. World Journal of Gastroenterology 15:1809-15.

Baeza AR. 2015. Roles of lipids and fatty acids through the spermatogenesis of European eel
Anguilla anguilla. MSc thesis, Universidad Polytechnic de Valencia.

Baris C. 2014. Fatty Acid Composition of the Muscle Lipids of Five Fish Species in Işıklı and
Karacaören Dam Lake. Veterinary Medicine International 2014: 5.

Cavington, MB. 2004. Omega-3 fatty acids. American Family Physician 70: 133-140.

Coetzee J, van der Lingen C, Hutchings L, Fairweather T. 2008. Has the fishery contributed to
a major shift in the distribution of South African Sardine? ICES: Journal of Marine Science
65: 1676–1688

Cook HW. 1996. Biochemistry of Lipids, Lipoproteins and Membranes. New Comprehensive
Biochemistry 36: 181-204.

Dalsgaard J, St John M, Kattner G, & Hagen W. 2003. Fatty acid trophic markers in the pelagic
marine environment. Advances in Marine Biology 46: 225-340.

Drevon CA. 2010. Fatty Acids: Structures and Properties. John Wiley & Sons Ltd.

Eschmeyer WN and Dempster LJ. 1990. Check-list of the fishes of the eastern tropical Atlantic
(CLOFETA). UNESCO 2: 665-679.

[11]
Field CJ. 2003. Fatty Acids dietary importance. Encyclopaedia of Food Sciences and Nutrition
(Second Edition). Academic Press Ltd.

Hamukuaya H, Bianchi G, and Baird D. 2001. The structure of demersal assemblages off
Namibia in relation to abiotic factors. South African Journal of Marine Science 23: 397-
417.

Hastings N, Agaba M, Tocher DR, Leaver MJ, Dick JR, Sargent JR, Teale AJ. 2001. A
vertebrate fatty acid desaturase with ∆5 and ∆6 activities. Proceedings of the National
Academy of Sciences 98: 14304–14309.

Heemstra PC. 1984. Family: Acropomatidae. FAO Species Identification Sheets for Fishery
Purposes. Western Indian Ocean Fishing Area 51: 8-12.

Iitembu JA, Richoux NB. 2016. Characterization of the dietary relationships of two sympatric
hake species, Merluccius capensis and M. paradoxus, in the northern Benguela region using
fatty acid profiles. African Journal of Marine Science 38: 39-48.

Iverson SJ, Frost KJ, and Lang SLC. 2002. Fat content and fatty acid composition of forage
fish and invertebrates in Prince William Sound, Alaska: Factors contributing to among and
within species variability. Marine Ecology Progress Series 241:161-181.

Izquierdo MS, Fernández-Palacios H & Tacon AGJ. 2001. Effect of brood stock nutrition on
reproductive performance of fish. Aquaculture 197: 25-42.

Jalaludin R. 2003. Analysis of fatty acids composition in marine fish (genus Tenualosa). MSc
thesis, Universiti Malaysia Sarawak.

Koop-man HN, Iverson SJ, and Gaskin DE. 1996. Stratification and age-related differences in
blubber fatty acids of the male harbour porpoise (Phocoena phocoena). Journal of
Comparative Physiology 165: 628–639.

Koussoroplis A, Bec A, Perga M, Koutrakis E, Desvilettes C, Bourdier G. 2010. Nutritional


importance of minor dietary sources for leaping grey mullet Liza saliens during settlement:
insights from fatty acid δ13C analysis. Marine Ecology Progress Series 404: 207–217.

[12]
Lambertsen G. 1977. Fatty Acid Compositions of Fish Fats. Comparisons Based on Eight Fatty
Acids. Journal of Arctic Biology 1: 105-116.

Logato PVR. 1998. Anatomia funcional e fisiologia dos peixes de água doce. Lavras. MG:
UFLA/FAEPE 108.

Macpherson E & Roel BA. 1987. Trophic relationships in the demersal fish community off
Namibia. South African Journal of Marine Science 5: 585-596.

Nichols DS. 2003. Prokaryotes and the input of polyunsaturated fatty acids to the marine food
web. FEMS Microbiology Letters 219: 1–7.

Palou A. Bonet ML. 2007. Novel Food Ingredients for Weight Control. Woodhead Publishing
Series. Retrieved from http://www.sciencedirect.com/science/book/9781845690304.

Robin JH, Regost C, Arzel J. & Kaushik SJ. 2003. Fatty acid profile of fish following a change
in dietary fatty acid source: model of fatty acid composition with a dilution hypothesis.
Aquaculture 225: 283 – 293.

Sara J, Iverson W & Chris F. 2012. Quantitative Fatty Acid Signature Analysis: A New Method
of Estimating Predator Diets. Ecological Monographs 74: 211-235.

Sakko AL. 1998. The influence of the Benguela upwelling system on Namibia’s marine
biodiversity. Biodiversity & Conservation 7: 419-433

Sargent JR, Bell JG, Bell MV, Henderson, RJ & Tocher DJ. 1993. The metabolism of
phospholipids and polyunsaturated fatty acids in fish. Aquaculture 43: 103–124.

Sargent JR, Henderson RJ & Tocher DR. 1989. The lipids. Fish Nutrition 2: 153- 218.

Sequeira V, Neves A, Vieira AR, Figueiredo I, Gordo LS. 2009. Age and growth of blue mouth,
Helicolenus dactylopterus, from the Portuguese continental slope. ICES Journal of Marine
Science 66: 524–531.

Steffens W. 1997. Effects of variation feeds on nutritive in essential fatty acids in fish value of
freshwater fish for humans. Aquaculture 151: 97-119.

[13]
Tocher DR and Ghioni M. 2010. Fatty acid requirements in ontogeny of marine and freshwater
fish. Aquaculture Research 41: 717-732.

Tocher DR. 2003. Metabolism and Functions of Lipids and Fatty Acids in Teleost Fish.
Reviews in Fisheries Science 11: 107–184.

Tulli F and Tibaldi E. 1997. Changes in amino acids and essential fatty acids during early larval
rearing of dentex. Aquaculture 5: 229-236.

[14]
6. Appendices
1. Fifty three (53) FAs ranging from 14 to 24 carbon atoms in length, were identified from
the samples.

Shorthand Common name Systematic name


14:0 Myristic acid Tetradecanoic acid
14:1
i-15:0
ai-15:0
15:0 Pentadecanoic acid
i-16:0
15:1
ai-16:0
16:0 Palmitic acid Hexadecanoic acid
16:1w7 Palmitoleic acid (Z) hexadec-9-enoic acid
16:1w5 Myristoleic acid c-9-tetradeconoic acid
i-17:0
ai-17:0
16:3w3 Hexadecatrienoic acid 7,10,13-hexacatrienoic acids
17:0 Margaric acid Heptadecanoic acid
16:3w4
17:1w7
16:4w3 4,7,10,13-hexadecatetraenoic
acid
16:4w1
18:0 Stearic acid Octadecanoic acid
18:1w9 Oleic acid (Z)-octadec-9-enoic acid
18:1w7 Vaccenic (Asclepic acid) 11-octadecenoic
18:1w6
18:1w5
18:2w6 Linoleic acid (9Z, 12Z)- octadeca-9,12-dienoic
acid
18:2w4
18:3w6 Gamma-linoleic acid (6Z,9Z,12Z)-octadeca-6,9,12-
trienoic acid
18:3w4
18:3w3 Alpha- linoleic acid (9Z,12Z,15Z)-octadeca-9,12,15-
trienoic acid
18:4w3 Stearidonic acid (6Z,9Z,12Z,15Z)-octadeca-
6,9,12,15-tetraenoic acid
18:4w1

[15]
20:0 Arachidic acid Icosanoic acid
20:1w9 Gondoic acid 11-eicosenoic acid
20:1w7
20:2w6
20:3w6 Dihomo-y-linoleic acid (8Z,11Z,14Z)-icosa-8,11,14-
trienoic acid
20:4w6 Arachidonic acid (5Z,8Z,11Z,14Z)-icosa-
5,8,11,14-tetraenoic acid
21:0
20:3w3 Eicosatrienoic acid 11,14,17-eicosatrienoic acid
20:4w3 Eicosatetraenoic acid 5,8,11,14,17-eicosatetraenoic
acid
20:5w3 Eicosapentaenoic acid (5Z,8Z,11Z,14Z,17Z)-icosa-
5,8,11,14,17-pentanoic acid
22:0 Behenic acid Docosanoic acid
22:1w11(13) Cetoleic acid 11-docosenoic acid
22:1w9 Erucic acid 13-docosenoic acid
21:5w3 Heneicosapentaenoic acid 6,9,12,15,18-
heneicosapentaenoic acid
22:2
23:0
22:4w6
22:4w3
22:5w6
22:5w3 Docosapentaenoic acid (7Z,10Z,13Z,16Z,19Z)-docosa-
7,10,13,16,19-pentaenoic aci
24:0 Lignoceric acid Tetracosanoic acid
22:6w3 Docosahexanoic acid (4Z,7Z,10Z,13Z,16Z,19Z,)-
docosa-4,7,10,13,16,19-
hexaenoic acid
24:6w3

2. Sampling Stations along the Namibian Atlantic ocean


Station no. Latitude Longitude Depth
3 -23,3433 13,48333 228
10 -24,3605 13,93067 244
13 -24,9337 13,96217 175
15 -25,805 14,20383 229
24 -27,1898 14,67333 274

[16]
25 -27,31 14,77833 287
28 -28,115 14,86333 197
37 -28,4762 14,34133 555
38 -27,5878 14,41333 411
40 -26,8402 14,38717 340
44 -25,873 13,766 393
45 -25,8413 13,55183 704

3. Species sampling stations and Lengths


Stations Species Length
3 Chlorophthamus agassizi 13
4 Chlorophthamus agassizi 13
4 Chlorophthamus agassizi 13
10 Chlorophthamus agassizi 15
10 Chlorophthamus agassizi 15
13 Chlorophthamus agassizi 15
13 Chlorophthamus agassizi 15
38 Chlorophthamus agassizi 15
4 Chlorophthamus agassizi 13
15 Helicolenus dactylopterus 12
15 Helicolenus dactylopterus 13
24 Helicolenus dactylopterus 18
25 Helicolenus dactylopterus 19
25 Helicolenus dactylopterus 20
28 Helicolenus dactylopterus 18
37 Helicolenus dactylopterus 14
44 Helicolenus dactylopterus 11
45 Helicolenus dactylopterus 15
45 Helicolenus dactylopterus 13
15 Helicolenus dactylopterus 13
28 Helicolenus dactylopterus 17
40 Synagrops microlepsis 12

[17]
40 Synagrops microlepsis 13
40 Synagrops microlepsis 14

4. Species Images

Jacopever (Helicolenus dactylopterus)

Shortnose greeneye (Chlorophthalmus agassizi)

Thinlip splitfin (Synagrops microlepsis)

[18]

You might also like