Isabel Ma ía Fe nández A iles
Deg ee in Ma ine Science (ULPGC)
isabel. e [email protected]
Academic yea 2013/2014
Tu o s: Ca men Ma ía He nández-C uz
Lidia Robaina Robaina
Final assignmen o he Ma ine Science
deg ee
E ec o di e en o i e s
en ichmen o mula eed on
g ow h and su i al o sea
b eam la ae (Spa us au a a)
Isabel Ma ía Fe nández A iles
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Index
Acknowledgemen s 3
TABLE LIST 4
FIGURE LIST 5
ABBREVIATIONS 6
1. In oduc ion 7
1.1 Global si ua ion o aquacul u e 7
1.2 La icul u e 8
1.3 Objec i es 11
2. Ma e ial and me hods 12
3. Resul s 15
3.1 G ow h 15
3.2 Su i al a e 17
3.3 Biomass 17
3.4 Ac i i y es 18
4. Discussion 19
5. Conclusions 22
6. Re e ences 23
Isabel Ma ía Fe nández A iles
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Acknowledgmen s
Desde es a página quisie a mos a mi ag adecimien o a odas aquellas pe sonas
que de una o ma u o a u ie on elación con la elabo ación de es e abajo. En
especial:
A mi amilia, sob e odo a mis abuelos, po su apoyo incondicional du an e oda la
ca e a y la ealización de es e abajo.
A la D a. Ca men Ma ía He nández-C uz, po su ines imable paciencia y po
ayuda me a es o za me cada día más.
A Lidia Robaina po la ayuda b indada, a pesa de que es u iese ocupada, siemp e
es u o ahí.
A Ángelo San ana, po su ayuda en los momen os más di íciles de es e abajo. Sin
él la es adís ica no se ía lo mismo.
A odos los écnicos del mesocosmos del Pa que cien í ico Tecnológico de Talia e,
po su acep ación como una más du an e los casi es meses que es u e allí.
A Desi, po su ines imable ayuda y paciencia a la ho a de lle a a cabo es e
expe imen o y po enseña me a con ia más en mí misma y se au osu icien e.
A Ma io, po su p edisposición a ayuda me, siemp e con una son isa.
A mis amigos y compañe os de clase que han es ado allí pa a da me ánimos en es a
úl ima e apa de la ca e a, especialmen e a Y onne, Dani y Jesús.
Isabel Ma ía Fe nández A iles
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TABLE LIST
Pages
Table I. La ae size, weigh , biomass, su i al a e and ac i i y es
along he expe imen . 15
Isabel Ma ía Fe nández A iles
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FIGURE LIST
Pages
Figu e 1. De elopmen o global aqua ic p oduc ion (aquacul u e and
ishe ies) om 1951 o 2011 7
Figu e 2. A e age size o 11 h DPH sea b eam la ae ed wi h he
di e en die s 16
Figu e 3. A e age size o 14 h DPH sea b eam la ae ed wi h he
di e en die s 16
Figu e 4. A e age weigh o 11 h DPH sea b eam la ae ed wi h he
di e en die s 16
Figu e 5. Su i al a e (%) o sea b eam la ae a he end o he
expe imen acco ding o di e en die s. The die s wi h he same le e s a e no
signi ican ly di e en (p- alue > 0.05) 17
Figu e 6. Biomass (µg) o sea b eam la ae a he end o he
expe imen acco ding o di e en die s 18
Figu e 7. La al su i al a e a e ac i i y es a he end o he
expe imen acco ding o di e en die s 18
Isabel Ma ía Fe nández A iles
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ABBREVIATIONS
FAO
APROMAR
PUFAs
EPA
DHA
UV
GIA
ANOVA
DPH
Food and Ag icul u e O ganiza ion o he Uni ed Na ions
Asociación Emp esa ial de P oduc o es de Cul i os Ma inos de España
Polyunsa u a ed a y acids
Eicosapen anoic acid 20:5 (n-3)
Docosahexanoic acid 22:6 (n-3)
Ul a iole
Aquacul u e Resea ch G oup
Analysis o a iance
Days pos ha ch
Isabel Ma ía Fe nández A iles
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1. In oduc ion
1.1 Global si ua ion o aquacul u e
Since he beginning o ci iliza ion, he ex ac ion o ma ine esou ces h ough
ishe ies and aquacul u e has been p esen . Some examples o his a e ound in Egyp o
China whe e, since ancien imes, ishes ha e been b eeding na u ally. E en now, you
can ind people aking ca e o ish in small pounds caused by he o e low o he Nile
Ri e in Egyp o in Chinese ice ields, whe e ca ps a e also a med wi h he ha es .
(Rueda, 2011)
Ac ually, he o igin o he ma ine esou ces ha we use is no only om ishe ies,
also aquacul u e plays a c ucial ole. Acco ding o epo s om he FAO in 2012, he
inc easing o ishe ies has made ha , abou 30% o he ish s ocks we e ound ully
exploi ed. Also, he consump ion pe capi a o sea ood (excluding algae) has inc eased
om 9.9 kg in 1960 o 19 kg in 2010. This has a o ed he boos o he aquacul u e in
he pas h ee decades, ha ing as a consequence an inc ease o he ish global
consump ion wi h 8.8% annual a e. This g ow h also is ela ed wi h an inc ease o i s
global ish con ibu ion om 9 - 49% in he las 30 yea s (APROMAR, 2013). Figu e 1.
Figu e 1. De elopmen o global aqua ic p oduc ion (aquacul u e and ishe ies) om 1951 o 2011.
APROMAR 2013
Cu en ly, conside ing only ish, ainbow ou (Onco hynchus mykiss), A lan ic
salmon (Salmo sala ) and sea b eam (Spa us au a a) a e, in ha o de , he ish species
widely cul u ed in Eu ope (APROMAR, 2013).
Isabel Ma ía Fe nández A iles
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1.2 La icul u e
Sea b eam (Spa us au a a) is one o he mos p oduced ish in Spain (APROMAR
2013). I is mainly due o i s ease o cul i a ion and also by i s high g ow h a e
(be ween 1.5 and 2 yea s o each comme cial size). Howe e , and despi e he high
game e p oduc ion o his specie du ing he h ee mon hs o b eeding, jus 90% o he
eggs a e e ilized and a ound 70% o hese ha ch success ully. Mo eo e , only among
20 - 35% o hese la ae pe cen age su i es (O ega, 2009). Mos o his high mo ali y
a e occu s du ing ansi ion among endogenous o exogenous eeding.
Acco ding o di e en au ho s (Pascual & Yú e a, 1987; Ri e a & Bo e o, 2009),
he mo ali y occu ence is mainly due o la ae mo phological limi a ion, in ol ing
mou h size, and also physiological es ic ions as a esul o an incomple e de elopmen
o he diges i e sys em. Th ough he yea s, a di ec ela ionship be ween he ype and
quali y o ood p o ided and he ex en o la al su i al has been demons a ed (Lazo,
2000; Conceição e al., 2010; Ham e e al., 2013).
The simplici y o diges i e ac in ha ched la ae, being li le mo e han a s aigh
ube wi hou di e en ia ion and wi h a e y low enzyme ac i i y (Go oni e al., 1986),
makes elabo a ed die s ine icien a his s age o de elopmen (Lazo, 2000; Izquie do e
al., 2000; Conceição e al., 2010).
Some au ho s such as Conceição e al. in 2010, no only poin ou o he diges i e
sys em o he la ae as he esponsible o a poo die . They also quali ies hem as isual
p eda o s, being he mo emen s o li ing p ey he igge o hei eeding esponse. This
p eda o y ins inc in he s ages o la al de elopmen suppo s zooplanck on li ing
p eys as he mos app op ia e die o la al eeding.
Acco ding o Ce ecedo-Ci e a e al. in 2004, o ensu e he supply o la ae by
li ing p ey; hey mus ha e ce ain equi emen s such as: ha ing he igh size o he
la ae mou h, a slow mo emen , a high a ailabili y and also, hey ha e o s imula e
la ae hun ing ins inc . Besides, he p eys mus each all minimum nu i ional
equi emen s and, he e o e, wi h easily diges ion o he la ae. Few species o
zooplank on each hese equi emen s; among hem we can ind o i e s (B achionus
sp.), b ine sh imp and copepods, being he i s wo species he mos widely cul i a ed
o he nu i ion pu pose (Lazo, 2000; Conceição e al., 2010).
Isabel Ma ía Fe nández A iles
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Some easons make o i e s an excellen sou ce o li ing p ey o la ae du ing i s
i s weeks o exogenous eeding: hey ha e a small size, be ween 70 - 350 μm, a high
g ow h a e, he capabili y o esis high popula ion densi ies and an ele a ed ole ance
ange o g owing condi ions and managemen . The o i e s also ha e an ele a ed
con en o p o ein, be ween 29 - 63% o i s d y weigh (Jeeja e al., 2011). Ne e heless,
only be ween 9 - 28% o i s d y weigh is lipid composi ion, which can a y acco ding
he die supplied o he o i e s (Conceição e al., 2010).
These lipids a e polyunsa u a ed a y acids (PUFAs), belonging o he g oup o
phospholipids, which se e as he main ene gy sou ce o ma ine ish la ae in hei
ea ly s ages (Rod iguez e al., 1997; Lazo, 2000; Bell e al., 2003; Ham e e al., 2013.).
Among hese a y acids, eicosapen aenoic acid (EPA) 20:5 (n-3) and docosahexaenoic
acid (DHA) 22:6 (n-3) plays a e y impo an ole. These PUFAs no only jus p o ide
ene gy o la ae; hey a e also essen ial o he cell memb anes in eg i y and unc ions
(Sa gen e al., 1997; Ri e a & Bo e o, 2009).
Despi e ha , some s udies show he limi ed capaci y o ish la ae o con e EPA
in o DHA. Sa gen e al. (1997) ha e demons a ed an impo an de iciency o he
enzyme del a-5-desa u ase, esponsible o enlonga ion and desa u a ion o PUFAs.
Besides, o i e s a e usually ed wi h bake 's yeas , which gi es hem a de icien
composi ion o PUFAs ha will be ansmi ed o he la ae by p ey inges ion. So, o
imp o e he up ake o hese a y acids, la ae a e eed wi h PUFAs high en iched
o i e s h ough mic oalgae o comme cial p oduc s (Ham e e al., 2008; Conceição e
al., 2010). This en ichmen mus ha e a 1:2 a io o EPA:DHA acco ding o Lazo
(2000), who p o ed ha , in excess, hese PUFAs gene a e ha m ul e ec s on neu onal
and op ical sys em o he la ae.
Along wi h a die ich in PUFAs, an app op ia e cul u e medium also p omo es
la al su i al. The echnique called "g een wa e ", which is based on he addi ion o
mic oalgae di ec ly o la al cul u e anks, has achie ed g ea esul s in e ms o
inc eased su i al, g ow h and inges ion a e o la ae (Øie e al., 1997; Rei an e al.,
1997). Rocha e al. (2008) also p o ed ha he p esence o mic oalgae in anks
inc eases he eeding esponse o la ae o sea b eam signi ican ly. Fu he mo e, he
“g een wa e ” echnique con ibu e o p ese e he nu i ional alue o p ey (Mak idis &
Isabel Ma ía Fe nández A iles
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0.0
0.5
1.0
1.5
2.0
2.5
3.0
3.5
4.0
4.5
5.0
5.5
6.0
Die 1
Die 2
Die 3
Die 4
Die 5
Size (mm)
DPH 11
Figu e 2. A e age size o 11 h DPH sea b eam la ae ed wi h he di e en die s.
0.0
0.5
1.0
1.5
2.0
2.5
3.0
3.5
4.0
4.5
5.0
5.5
6.0
6.5
Die 1
Die 2
Die 3
Die 4
Die 5
Size (mm)
DPH 14
Figu e 3. A e age size o 14 h DPH sea b eam la ae ed wi h he di e en die s.
0.00
0.05
0.10
0.15
0.20
0.25
0.30
0.35
Die 1
Die 2
Die 3
Die 4
Die 5
Weigh (μg)
DPH 11
Figu e 4. A e age weigh o 11 h DPH sea b eam la ae ed wi h he di e en die s.
.
Isabel Ma ía Fe nández A iles
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3.2 Su i al a e
Rega ding su i al a es, he da a ob ained a he end o he expe imen showed
signi ican di e ences a e pe o m s a is ical analysis (p- alue = 0.013). These
di e ences we e ound be ween die 3 (Isoch ysis sp.), which had he highes su i al
a e, and die s 1 (DHA P o ein SELCO) and 4 (Dunaliella sp.), bo h wi h he lowes
su i al a e as is shown in igu e 5.
Howe e , due o he abno mali y o he die 1 da a, which should ha e been one o
he bes su i al a es, i is hough ha i could be a p oblem wi h he p oduc .
0.0
1.0
2.0
3.0
4.0
5.0
6.0
7.0
8.0
Die 1
Die 2
Die 3
Die 4
Die 5
Su i al a e (%)
Figu e 5. Su i al a e (%) o sea b eam la ae a he end o he expe imen acco ding o di e en die s.
The die s wi h he same le e s a e no signi ican ly di e en (p- alue > 0.05)
3.3 Biomass
Once he expe imen ended, all he anks had a a iable numbe o su i al la ae
o he coun o he biomass and i s subsequen s a is ical analysis. As we can see in he
igu e 6, i s dis ibu ion is simila o he ob ained o su i al a e excep ha , in his
case, he di e ences o la ae biomass ound be ween die s we e no signi ican .
I was also shown ha he highes biomass was ound in he anks whe e la ae
we e ed wi h die 3 (167.57 mg) while he lowes we e again in anks whose la ae
we e ed wi h die s 1 (DHA P o ein SELCO) and 4 (Dunaliella sp.), being he las one
he die wi h he lowes ob ained biomass (Figu e 6).
a
a
b
ab
ab
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0
20
40
60
80
100
120
140
160
180
Die 1
Die 2
Die 3
Die 4
Die 5
Biomass (μg)
Figu e 6. Biomass (µg) o sea b eam la ae a he end o he expe imen acco ding o di e en die s.
3.4 Ac i i y es
Obse ing he esul s o ac i i y es (Figu e 7), a simila dis ibu ion was ound o
hose ob ained o su i al and biomass (Figu es 5 and 6 espec i ely). I was mo eo e
obse ed ha la ae ed wi h die s 1 and 4 showed a dec easing su i al a e a e he
applied s ess, whe eas la ae ed wi h die 3 we e he mos a e mo e esis an , wi h
86.67% o su i al.
0
10
20
30
40
50
60
70
80
90
100
Die 1
Die 2
Die 3
Die 4
Die 5
Su i al a e (%)
Figu e 7. La al su i al a e a e ac i i y es a he end o he expe imen acco ding o di e en die s.
Isabel Ma ía Fe nández A iles
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4. Discussion
Analyzing he esul s, we can’ see any concluding esponse be ween es ed die s
ega ding he size o sea b eam la ae a he samplings o he expe imen . Bu wi h
weigh da a ob ained, we can see a sligh end owa d highe weigh in die 5 (bake ’s
yeas ). Tha end migh be ela ed wi h low biomass le els ound in he la ae anks ed
wi h ha die .
This inc ease in la al g ow h associa ed wi h dec eased biomass has also been
desc ibed by Roo e al. in 2010. They es ed la al g ow h a di e en densi ies,
esul ing in a highe g ow h a e a low densi ies (> 50 la ae L-1). So, hey p o ed ha
high la al densi ies a e associa ed wi h low a es o g ow h due o a dec eased appe i e,
an inc eased agg essi eness o la ae and he e o e an in ense in aspeci ic compe i ion
o ood esou ces. All his igge ed a ch onic s ess si ua ion in la ae, dec easing he
biomass o he sys em and a o ing he su i al o la ae be e adap ed and mo e
esis an (He nandez-C uz e al., 1999).
Howe e , he abo e desc ibed o biomass o die s 1 (DHA P o ein SELCO) and 4
(Dunaliella sp.), whe e he e a e e en less biomass in he anks han in die 5, does no
co espond wi h he esul s ob ained o a e age weigh , which alues should be highe .
Du ing his wo k, i has been exposed a many imes he g ea impo ance o he
in ake o hese a y acids in he la ae. Many au ho s ha e ound in hem he esponse
o an inc ease o dec ease o g ow h and su i al in hese ea ly s ages o la al
de elopmen (Salhi, 1997; Rod íguez e al., 1994; Salhi e al., 1994). The da a ob ained
o die s 1 and 4, does no i wi h he desc ibed by He nández-C uz e al. (1999) and
Roo e al. (2010), showing hen a poo ly PUFAs en iched die o la ae.
Yeas supplied o o i e s in he die 5, had a ound 1.3% o PUFAs [1] while he
die 4, made om he mic oalga Dunaliella sp., has only a ound 0.4 - 0.8% o hese
a y acids (Ne ejan e al., 2003). This ag ees wi h he da a obse ed o us.
Howe e , he da a ob ained o die 1 do no co espond o expec a ions. A s udy by
Sil a in 1999 e alua ed he con en o PUFAs in o i e s en iched wi h DHA P o ein
SELCO, ob aining le els o EPA and DHA o 12.1 and 25.2 mg g-1 o d y weigh o
EPA and DHA espec i ely.
Isabel Ma ía Fe nández A iles
20
These le els canno be compa ed wi h hose ob ained o o i e s en iched wi h
yeas which, acco ding o Wa anabe e al. in 1983, anging om 1 o 2 mg g-1 o d y
weigh o EPA. This is why i ’s hough ha he anomalous da a ound in he die 1
could be explained as a possible esul o a a y acid oxida ion, pe haps due o a poo
main enance o he p oduc . This would be cla i ied in pa analyzing he biochemical
composi ion o he die s, bu ha is no he objec i e o his expe imen .
On he o he hand, we also ha e high le els o PUFAs in die 3, made om
Isoch ysis sp., which do no show an inc ease in a e age la al g ow h compa ed wi h
he o he die s es ed wi h lowe con en o hese a y acids. Acco ding Hue limann e
al. (2010), his mic oalgae has a ound 8.5 - 15.9% o d y weigh o PUFAs. Sil a in
1999 pe o med an expe imen o analyze he le el o PUFAs in Isoch ysis galbana
en iched o i e s, ob aining a composi ion be ween 15 - 20 mg g-1 o d y weigh in EPA
and DHA. These alues a e e y simila o hose ob ained by his au ho in o i e s
en iched wi h DHA P o ein SELCO.
Howe e , o he anks ed wi h die 3, high biomass was obse ed, which we e
also epo ed by He nandez-C uz e al. (1999), and la e on by Roo e al. (2010). They
s a ed ha an inc ease biomass is associa ed wi h educed g ow h o he la ae.
Howe e , nei he he su i al a e no he la ae esis ance we e no educed by his
high biomass due o he high PUFAs le els in he die .
Simila ly wi h he die 3, he die 2 (Te aselmis sp.) exhibi s low alues o a e age
weigh and high in e ms o biomass, su i al and esis ance. Howe e , despi e ha ing
ela i ely high le els o PUFAs (0.7 - 3.8% o d y weigh o algae acco ding
Hue limann e al. in 2010), his die does no exceed he alues achie ed by die 3 in
any espec .
Se e al au ho s a gue ha his may be due o he high equi emen s o DHA
compa ed o EPA in la ae as a esul o he high a e o g ow h and de elopmen o
issues and s uc u es in he la al s age (Takeuchi e al., 1994; Ri e a & Bo e o, 2009).
DHA has been desc ibed as an impo an componen o he issue neu al la ae and also
imp o es ision and, consequen ly, be e ca ch p ey which leads o a be e ed
(Rod iguez e al., 1997; Fu ui a e al., 2000; Ri e a & Bo e o, 2009). Tha is he eason
why, despi e ha die 3 has a lowe con en o EPA han die 2, possessing a much
highe alue o he DHA imp o es la ae de elopmen .
Isabel Ma ía Fe nández A iles
21
Finally, la ae ed wi h die s 2 and 3 showed a highe esis ance o s ess caused by
he ac i i y es , bu la ae ed wi h die 3 ob ained he bes esul s. An expe imen
conduc ed by Takeuchi e al. (1991) wi h la ae o Japanese sea b eam (Pag us majo ),
showed ha hose ed la ae wi h a DHA ich die s pe o med be e su i al a e be o e
he es ac i i y han whose la ae which we e ed wi h die s ich in EPA.
Isabel Ma ía Fe nández A iles
22
5. Conclusions
The g ow h o he la ae is highly dependen on he ank la ae densi y, and hus
highe in ensi y o la ae compe i ion du ing eeding. A educ ion o numbe o la ae
pe ank appea s a possible educ ion o his p oblem.
Among he es ed die s, he use o mic oalgae Dunaliella sp. o o i e s en ichmen
does no p o ide accep able esul s in la al g ow h and su i al a e. By he o he
hand, he mic oalgae Te aselmis sp., and despi e i s high le els o EPA, does no
p o ide enough PUFAs o la ae mainly by i s low DHA con en .
High DHA con en die s, as ha con aining Isoc hysis sp. in p esen expe imen ,
p omo e be e su i al and s ess esis ance o he la ae.
Isabel Ma ía Fe nández A iles
23
6. Re e ences
APROMAR (2013). La acuicul u a en España 2013. APROMAR, ESACUA, Fundación
OESA
Bell, J. G., McE oy, L. A., Es e ez, A., Shields, R. J., & Sa gen , J. R. (2003).
Op imising lipid nu i ion in i s - eeding la ish la ae. Aquacul u e, 227(1),
211-220.
Ci e a-Ce ecedo, R., Al a ez-González, C. A., & Moyano-López, F. J. (2004).
Nu ición y alimen ación de la as de peces ma inos. A ances en nu ición
acuícola VII. Memo ias del VII Simposium In e nacional de nu ición Acuícola,
He mosillo, Sono a, México.
FAO (2012). El es ado mundial de la pesca y la acuicul u a 2012. Roma. 231 págs.
Fu ui a, H., Tanaka, H., Yamamo o, T., Shi aishi, M., & Takeuchi, T. (2000). E ec s o
n-3 HUFA le els in b oods ock die on he ep oduc i e pe o mance and egg and
la al quali y o he Japanese lounde , Pa alich hys oli aceus. Aquacul u e,
187(3), 387-398.
Go oni, J. J., Boehle , G. W., & Wa anabe, Y. (1986). The physiology o diges ion in
ish la ae. En i onmen al Biology o Fishes, 16(1-3), 59-77
Ham e, K., S i as a a, A., Rønnes ad, I., Mango ‐Jensen, A., & S oss, J. (2008).
Se e al mic onu ien s in he o i e B achionus sp. may no ul il he nu i ional
equi emen s o ma ine ish la ae. Aquacul u e Nu i ion, 14(1), 51-60.
Ham e, K., Yú e a, M., Rønnes ad, I., Boglione, C., Conceição, L. E., & Izquie do, M.
(2013). Fish la al nu i ion and eed o mula ion: knowledge gaps and
bo lenecks o ad ances in la al ea ing. Re iews in Aquacul u e, 5(s1), S26-
S58.
He nández-C uz, C. M., Salhi, M., Bessona , M., Izquie do, M. S., González, M. M., &
Fe nández-Palacios, H. (1999). Rea ing echniques o ed po gy (Pag us pag us)
du ing la al de elopmen . Aquacul u e, 179(1), 489-497.
Hue limann, R., De Nys, R., & Heimann, K. (2010). G ow h, lipid con en ,
p oduc i i y, and a y acid composi ion o opical mic oalgae o scale‐up
p oduc ion. Bio echnology and bioenginee ing, 107(2), 245-257.
Izquie do, M. S., J. Soco o, L. A an zamendi y C. M. He nández-C uz. (2000).
Diges ión, abso ción y u ilización de lípidos en la as de peces ma inos. pp 251-
263. A ances en Nu ición Acuícola IV. Memo ias del IV Simposium
In e nacional de Nu ición Acuícola. La Paz, B.C.S., México.
Isabel Ma ía Fe nández A iles
24
Jeeja, P. K., Imelda, J., & Paul aj, R. (2011). Nu i ional composi ion o o i e
(B achionus plica ilis Mulle ) cul u ed using selec ed na u al die s. Indian Jou nal
o Fishe ies, 58(2), 59-65.
Lazo, J. (2000). Conocimien o ac ual y nue as pe spec i as en el desa ollo de die as
pa a la as de peces ma inos. A ances en Nu ición Acuícola V. Memo ias del V
Simposium In e nacional de Nu ición Acuícola. Mé ida, Yuca án, Mexico.
Mak idis, P., & Vads ein, O. (1999). Food size selec i i y o A emia anciscana a
h ee de elopmen al s ages. Jou nal o plank on esea ch, 21(11), 2191-2201.
Naas, K. E., & Ha boe, T. (1992). Enhanced i s eeding o halibu la ae
(Hippoglossus hippoglossus L.) in g een wa e . Aquacul u e, 105(2), 143-156.
Naas, K., Huse, I., & Iglesias, J. (1996). Illumina ion in i s eeding anks o ma ine
ish la ae. Aquacul u al Enginee ing, 15(4), 291-300.
Ne ejan N., Saez I., Gaja do G., So geloos P. (2003) Supplemen a ion o EPA and
DHA emulsions o a Dunaliella e iolec a die : e ec on g ow h and lipid
composi ion o scallop la ae, A gopec en pu pu a us (Lama ck, 1819).
Aquacul u e, 217 pp. 613–632
Øie, G., Mak idis, P., Rei an, K. I., & Olsen, Y. (1997). P o ein and ca bon u iliza ion
o o i e s (B achionus plica ilis) in i s eeding o u bo la ae (Scoph halmus
maximus L.). Aquacul u e, 153(1), 103-122.
O ega G. A. (2009). Cul i o de Do ada (Spa us au a a). Mad id: Fundacin
Obse a o io de Acuicul u a.
Pascual, E., & Yú e a, M. (1987). Alimen ación en el cul i o la a io de peces ma inos.
Rei an, K. I., Rainuzzo, J. R., Øie, G., & Olsen, Y. (1997). A e iew o he nu i ional
e ec s o algae in ma ine ish la ae. Aquacul u e, 155(1), 207-221.
Ri e a C. M., Bo e o Z. M. (2009) Alimen o i o en iquecido con ácidos g asos pa a el
desa ollo la a io de peces. Re Colomb Cienc Pecu 2009; 22: 607-618
Rocha, R. J., Ribei o, L., Cos a, R., & Dinis, M. T. (2008). Does he p esence o
mic oalgae in luence ish la ae p ey cap u e?. Aquacul u e esea ch, 39(4), 362-
369.
Rod iguez, C., Pe ez, J. A., Izquie do, M. S., Mo a, J., Lo enzo, A., &
Fe nandez‐Palacios, H. (1994). Essen ial a y acid equi emen s o la al gil head
sea b eam, Spa us au a a (L.). Aquacul u e Resea ch, 25(3), 295-304.
Rod iguez, C., Pe ez, J. A., Diaz, M., Izquie do, M. S., Fe nández-Palacios, H., &
Lo enzo, A. (1997). In luence o he EPA/DHA a io in o i e s on gil head
seab eam (Spa us au a a) la al de elopmen . Aquacul u e, 150(1), 77-89.
Isabel Ma ía Fe nández A iles
25
Roo, J., He nández-C uz, C. M., Bo e o, C., Schucha d , D., & Fe nández-Palacios, H.
(2010). E ec o la al densi y and eeding sequence on meag e (A gy osomus
egius; Asso, 1801) la al ea ing. Aquacul u e, 302(1), 82-88.
Rueda F. M (2011). B e e his o ia de una g an desconocida: la acuicul u a. Re is a
Eubac e ia Nº 26 No iemb e 2011 h p://www.um.es/eubac e ia/acuicul u a.pd
Salhi, M., Izquie do, M. S., He nandez-C uz, C. M., Gonzalez, M., & Fe nandez-
Palacios, H. (1994). E ec o lipid and n−3 HUFA le els in mic odie s on
g ow h, su i al and a y acid composi ion o la al gil head seab eam (Spa us
au a a). Aquacul u e, 124(1), 275-282.
Salhi, M. (1997). Es udio de los eque imien os lipídicos de la as de Do ada (Spa us
au a a) alimen adas con mic odie as. Tesis Doc o al. Uni e sidad de Las Palmas
de G an Cana ia, España, 178 pp.
Sa gen , J. R., McE oy, L. A., & Bell, J. G. (1997). Requi emen s, p esen a ion and
sou ces o polyunsa u a ed a y acids in ma ine ish la al eeds. Aquacul u e,
155(1), 117-127.
Sil a, A. (1999). E ec o de la mic oalga Isoch ysis galbana en el cul i o emp ano de
la as de Pa alich hys adspe sus. Ciencias Ma inas, 25(2).
Takeuchi T, Toyo a M, Wa anabe T. (1991) Die a y alue o ed seab eam o A emia
nauplii en iched wi h EPA and DHA. Abs ac s o he Annual Mee ing o
Japanese Socie y o Scien i ic Fishe ies, Tokyo; p. 243.
Takeuchi T, Feng Z, Yoseda K, Hi okawa J, Wa anabe T. (1994). Nu i i e alue o
DHA-en iched o la al cod. Nippon Suisan Gakkaishi; 60:641-652.
Wa anabe, T., Ki ajima, C., & Fuji a, S. (1983). Nu i ional alues o li e o ganisms
used in Japan o mass p opaga ion o ish: a e iew. Aquacul u e, 34(1), 115-143.
[1] h p://www. ao.o g/doc ep/ ield/003/ab473s/AB473S03.h m#chIII