EMBRYONIC DEVELOPMENT AND NUTRITIONAL REQUIREMENTS OF KUTUM FRY, RUTILUS FRISII KUTUM

DAVOUD TALEBI HAGHIGHI

DOCTOR OF PHILOSOPHY UNIVERSITI PUTRA MALAYSIA

2006

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EMBRYONIC DEVELOPMENT AND NUTRITIONAL REQUIREMENTS OF KUTUM FRY, RUTILUS FRISII KUTUM

BY

DAVOUD TALEBI HAGHIGHI

Thesis Submitted to the School of Gaduate Studies, Universiti Putra Malaysia, in Fulfilment of the Requirement for the Degree of Doctor of Philosophy

August 2006

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DEDICATION

To my parents

For their true love, constant trust and helps

To my wife

For her kindness, devotion and support during all difficulties

To my children

For their undetstanding, dignities and tolerance

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Abstract of thesis presented to the Senate of Universiti Putra Malaysia in fulfilment of the requirement for the degree of Doctor of Philosophy

EMBRYONIC DEVELOPMENT AND NUTRITIONAL REQUIREMENTS OF KUTUM FRY, RUTILUS FRISII KUTUM

By

DAVOUD TALEBI HAGHIGHI

August 2006

Chairman: Associate Professor Che Roos Saad, PhD

Faculty: Agriculture

This study consisted of a series of experiments that were conducted to

determine some biological characteristics, artificial reproduction, embryonic

development, optimal level of protein and lipid in artificial diets and food regime

spectrum in the stage of secondary nursing of kutum fish larvae (Rutilus frisii

kutum).

The above study was carried out with artificial reproduction to determine some

biological characteristics such as: age, weight, length, percentages of ovary

weight to broodstock weight, and also estimation of absolute and relative

fecundity and evaluated their relationships in kutum fish broodstock. The results

of this study showed that the optimal ratio of female to male was 1:2 (♀ :♂), the

first sexual maturity occurred at 2 year old with well developed sexual products

with length ranging from 25 cm to 48cm and weight ranging from 213.5 g to 1588

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g. Most age, length and weight–frequencies belonged to age groups III+

(23.20%) to IV+ (20%). The results of this study indicated that the absolute

fecundity increased as the fish grows, ranging from 12022 to 78856 eggs per

fish; on the contrary, relative fecundity of fish decreased with the increasing of

fish age and it ranged from 67 to 42 egg for per g of ovary for age groups II+ to

VII+ respectively. Correlation coefficient for absolute fecundity against length,

weight and age were observed to be r=0.76, r=0.68 and r=0.73 respectively, but

the correlation coefficient between relative fecundity against length, weight and

age were lower with values of r=0.0.37, r=0.30 and r=0.33 respectively.

The stripping of eggs was performed in semi-dry condition. Males and females

were injected with carp pituitary gland extracts (GnRh) to induce ovulation at 2

mg of GnRh per kilogram of broodstock body weight. Seven period of

embryogenesis were defined; I) the newly fertilized egg was zygote, occurred

with 1 cell, with an average egg diameter of 2.21 mm, and mean weight of 9.99

mg and lasted for 15 min. II) Cleavage stage began from 2 to 64 divided cells,

with egg diameter of 2.715 mm, weight at 12.99 mg, in 7 hours. III) Blastula

stage was made up of early epiboly at the animal pole, with egg diameter 2.8

mm, with weight of 13.51 mg, and time of 26 hours. IV) Gastrula stage was

formed when two layers of cells including epiblast which gave rise to ectoderm

tissues and hypoblast which gave rise to both mesoderm and endoderm;

consequently, the embryonic axis and epiboly were formed at the 90% of

gastrulation period. It lasted for 33 h, with egg diameter of 2.96 mm and egg

weight of 13.61 mg. V) Segmentation stage began with the formation of the

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primary organs; somites, early organogenesis, neuromeres, elongation of tail

and earliest movement appeared. It lasted for 29 h, with diameter of egg at 3.04

mm, weight egg of 13.24 mg. VI) Organogenesis occurred with rapid

development towards functionality in most organs, particularly the senses, the

heart and the gut. It lasted for 15 h with egg diameter of 3.16 mm and weight of

14.64 mg. VII) Hatching ended when the larvae were pelagic and avoids sinking.

It lasted 20 h after organogenesis with diameter egg 3.009 mm and weight 14.39

mg. Total times for hatching was 130 hours and 15 minutes.

Food regime spectrum of kutum fry at the secondary nursing under pond culture

condition was estimated based on their stomach contents analysis. During the

rearing period water temperature varied from 16 °C to 28 °C. In general, the

frequency and diversity of phytoplankton included 5 phylums, 10 orders, 21

families and 36 genuses and zooplankton community included 5 phylums, 10

orders, 12 families and 31 genuses were obtained from gut contents analysis of

kutum fry. Observation was carried out

Observation was carried out periodically for 11 weeks, and it was found that

kutum fry collected its food from below to surface of water. The results of this

study indicated that kutum fry had low growth and high mortality at the early

stage of rearing period. When the intestinal digestion system became more

developed, the utilization of greater diversity of food items increased. The

feeding regime spectrum of kutum fry consisted of phytoplankton, zooplankton,

benthos, detritus and artificial food.

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Determination of optimal level of crude protein in kutum fry diet was conducted

with five levels of protein concentrations (35%, 40%, 45%, 50% and 55%). The

results showed significant difference among 5 concentrations on growth of fish

(p<0.05). The best final weight was 1095 mg, weight gain of 877.3%, specific

growth rate of 3.21%, protein efficiency ratio of 1.00, food conversion ratio of

1.99, and at 28% mortality obtained at 50% protein level diet. Polynomial

regression analysis between protein levels and weight gain of fish indicated that

a weight gain peak was obtained at 46.8% protein level. Results of this

experiment showed that the optimum protein level of kutum larvae for best

growth was 46.8%.

Determination of optimal level of crude lipid in kutum fish fry diet was conducted

with four levels of lipid concentrations (8%, 10%, 12% and 14%). The results

showed significant difference among lipid levels on growth of fish (P< 0.05). The

highest final weight was 1926 mg, weight gain 852%, specific growth rate 2.78%,

protein efficiency ratio 1.07, food conversion ratio 2.05 and mortality 2.5%

obtained when dietary lipid level was 8%. Polynomial

regression analysis between lipid levels and weight gain of fish indicated that a

weight gain peak was at 7.67% lipid level. Result of this experiment showed that

the optimum level lipid of kutum larvae for best growth was at 7.67% lipid level.

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Abstrak tesis yang dikemukakan kepada Senat Universiti Putra Malaysia sebagai memenuhi keperluan untuk ijazah Doktor Falsafah

PERKEMBANGAN EMBRIO DAN KEPERLUAN NUTRIEN FRI KUTUM, RUTILUS FRISII KUTUM

Oleh

DAVOUD TALEBI HAGHIGHI

August 2006

Pengerusi: Profesor Madya Che Roos Saad, PhD

Fakulti: Pertanian

Satu siri eksperimen telah dijalankan untuk menentukan beberapa ciri biologi,

pembiakan aruhan, perkembangan embrio, paras protein dan lipid yang

optimum dalam diet buatan dan spektrum makanan pada peringkat penjagaan

sekunder bagi fri ikan kutum (Rutilus frisii kutum).

Kajian di atas telah dijalankan menggunakan pembiakan aruhan untuk

menentukan ciri-ciri biologikal seperti: umur, berat, panjang kelas, peratus berat

ovari kepada berat ikan induk, anggaran fekunditi absolut dan fekunditi relatif

serta menilai perhubungan tersebut terhadap induk ikan kutum. Hasil kajian

menunjukkan bahawa nisbah optimum induk spesis jantan kepada betina adalah

sama 1:2 (♀:♂); kematangan seksual paling awal berlaku pada umur 2 tahun

dengan pembentukan produk seksual yang matang sehingga umur 7 tahun,

dengan panjang ikan dalam lingkungan 25cm ke 48cm dan berat

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daripada 213.5 g ke 1588 g. Kebanyakan umur, panjang dan kekerapan berat

tergolong dalam kumpulan umur III+ – IV+. Hasil kajian ini juga menyatakan

bahawa fekunditi keseluruhan semakin meningkat mengikut pembesaran ikan

iaitu daripada 12022 kepada 78856 bilangan telur bagi setiap ikan atau ovari.

Walau bagaimanapun, fekunditi relatif ikan semakin berkurang dengan

peningkatan umur ikan dan termasuk dalam lingkungan 67 ke 42 telur sehari

bagi setiap gram ovari bagi kumpulan umur tersebut, masing-masing dalam

lingkungan II+ ke VII+ setiap satu. Pekali koefisien korelasi antara fekunditi

keseluruhan terhadap panjang, berat dan umur adalah r=0.76, r=0.68 dan r=0.73

masing-masing, tetapi pekali korelasi antara fekunditi relatif terhadap panjang,

berat dan umur adalah lebih rendah iaitu r=0.37, r=0.30 and r=0.33 masing-

masing.

Proses melurut telur ikan telah dijalankan dalam keadaan separa kering. Ikan

jantan dan betina telah disuntik dengan ekstrak daripada kelenjar pituitari

sebanyak 2 mg GnRh setiap kilogram berat ikan untuk mempercepatkan

kematangan. Sebanyak 7 peringkat pembentukan embrio yang terlibat telah

didefinasikan sebagai: I) Zigot, telur baru yang telah disuburkan, terbentuk

dengan 1 sel dan diameter purata telur adalah 2.21 mm, min berat sebanyak

9.99 mg dan mengambil masa sehingga 15 min. II) Peringkat Cleavage bermula

daripada pembahagian sel daripada 2 hingga 64, dengan diameter telur

sebanyak 2.715 mm, berat sebanyak 12.99 mg, dalam masa 6 jam. III)

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Peringkat Blastula terdiri daripada epiboli awal pada kutub haiwan, dengan

diameter telur sebanyak 2.8 mm, berat sebanyak 13.51 mg, selama 26 jam. IV)

Peringkat Gastrula pula terbentuk apabila 2 lapisan sel termasuklah epiblast

yang telah bertukar kepada tisu ektoderma dan hipoblast yang juga telah

bertukar kepada tisu mesoderma dan endoderma; pada masa yang sama paksi

embrionik telah dibentuk dan epiboli terbentuk pada 90% proses grastulasi. Ia

berlangsung selama 33 jam, dengan diameter telur sebanyak 2.96 mm dan berat

telur sebanyak 13.61 mg. V) Peringkat Segmentasi bermula dengan

pembentukan organ-organ primer, somit, organogenesis awal, neuromeres,

pemanjangan ekor dan berlakunya pergerakan awal. Ia berakhir dalam masa 29

jam, dengan diameter dan berat telur sebanyak 3.04 mm dan 13.24 mg masing-

masing. VI) Organogenesis berlaku dengan perkembangan yang pantas ke

arah kefungsian sebahagian besar organ, terutamanya organ sensori, jantung

dan gastrousus. Ia berakhir selama 15 jam dengan diameter dan berat telur

sebanyak 3.16 mm dan 14.64 mg. VII) Penetasan berakhir apabila larva bersifat

pelagik dan dapat mengelak dari tenggelam. Proses ini berlaku dalam tempoh

20 jam selepas organogenesis dengan diameter telur sebanyak 3.009 mm dan

berat sebanyak 14.39 mg. Tempoh penetasan keseluruhan adalah selama 129

jam 25 min. Spektrum makanan fri ikan kutum pada penjagaan sekunder di

bawah keadaan kultur kolam telah dianggarkan berdasarkan analisis kandungan

perut ikan. Semasa tempoh pemeliharaan, suhu air dan keadaan cuaca berbeza

antara 16°C dan 28°C. Secara amnya, kekerapan dan kepelbagaian fitoplankton

termasuklah 5 filum, 10 order, 21 famili dan 36 genus sementara komuniti

zooplankton pula terdiri daripada 5 filum, 10 order, 12 famili dan 31 genus yang

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diperolehi daripada analisis kandungan perut fri ikan kutum. Pemerhatian telah

dijalankan secara berkala setiap 2 minggu selama 11 minggu, dan didapati

bahawa fri ikan kutum mendapatkan makanan dari bahagian bawah sehinggalah

ke permukaan air. Spektrum rejim pemakanan bagi fri ikan kutum terdiri daripada

fitoplankton, zooplankton, benthos, detritus dan makanan tiruan. Hasil kajian

mendapati bahawa fri ikan kutum mempunyai kadar tumbesaran yang rendah

dan kadar kematian yang tinggi pada peringkat awal tempoh penternakan.

Apabila sistem penghadaman usus terbentuk, penggunaan jenis-jenis makanan

yang berbeza dan pelbagai dipertingkatkan.

Penentuan paras optimum kandungan protein kasar dalam diet fri ikan kutum

telah dijalankan mengikut lima kepekatan protein (35%, 40%, 45%, 50%, 55%)

yang menunjukkan perbezaan yang signifikan terhadap pertumbuhan ikan (P<

0.05). Nilai terbaik bagi berat akhir, pertambahan berat, kadar pertumbuhan

spesifik, nisbah kecekapan protein, nisbah pertukaran makanan dan kematian

adalah 1095 mg, 877.3 %, 3.21 %, 1.00 %, 1.99 dan 18 % masing-masing

apabila diberi diet yang mengandungi kandungan protein sebanyak 50 %.

Analisis regresi polinomial di antara paras protein dan pertambahan berat badan

ikan menunjukkan bahawa puncak pertambahan berat ikan adalah pada paras

protein 46.8%. Keputusan eksperimen ini menunjukkan bahawa paras protein

diet yang optimum untuk pertumbuhan larva ikan kutum yang terbaik adalah

pada 46.8%.

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Penentuan paras optimum kandungan lemak kasar dalam diet fri ikan kutum

telah dijalankan mengikut empat kumpulan dengan kepekatan (8%, 10%, 12%,

14%) yang menunjukkan perbezaan yang signifikan terhadap pertumbuhan ikan

(P< 0.05). Nilai tertinggi bagi berat akhir, pertambahan berat, kadar pertumbuhan

spesifik, nisbah keberkesanan protein, nisbah pertukaran makanan dan

kematian adalah 1926 mg, 852%, 2.78%, 1.07, 2.05 dan 2.5% masing, yang

diperolehi pada paras lipid diet adalah 8 %. Analisis Regresi Polinomial di antara

paras lipid dan pertambahan berat badan ikan menunjukkan bahawa puncak

pertambahan berat ikan adalah pada paras lipid 7.67%. Keputusan eksperimen

ini menunjukkan bahawa paras lipid diet yang optimum untuk pertumbuhan larva

ikan kutum yang terbaik adalah pada 7.67%.

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ACKNOWLEDGEMENTS In the name of GOD

I would like to express my sincere gratitude and thank to the chairman of my

Supervisor Committee, Associate Professor Dr. Che Roos Saad, for his support

and valuable guidance throughout my doctoral study.

My sincerest appreciation is also extended to Prof. Dr. Abul Razak Alimon who

is a member of my Supervisory Committee, for his worthful guidance and

constructive suggestions during the study period.

I would also like to thank Prof. Dr. Ali Nikkhah, another member of my

Supervisory Committee, for his help as well as guidance during the entire study

period.

I wish to express my thanks to Dr. Rezvani Director of Fisheries Research and

Training Organization, and Dr. Khanipour Head of Gillan Fisheries Research

Center, without whose understanding, help and support this research could not

have been realized.

Thanks are extended to Frshad Mahisfat, Shafizadeh, Afshin Amiri, Alireza

Afraz, Mahdizadeh, Moradi, Mkaremi and Saffai Head of Ghazian Station and

also staff of Chemistry Department of Gillan Fishery Research, especially

Tajadod.

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I would like to thank from all the staff in Shahid Anssari Reproduction Complex,

especially Mr. Mohamd Tolloi and Mr. Khomairani.

I am also very thankful to the staff in Food Analysis Laboratory of Gillan

Province Veterinary Department, Dr. Zahdi, Mr. Taghavi and Mr. Ghasami.

Finally, my deep appreciation goes to my wife, Dr. Saydeh Zahra Khatami, my

children and my entire family for their support and encouragement.

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I certify that an Examination Committee has met on 9th August 2006 to conduct the final examination of Davoud Talebi Haghighi on his Doctor of Philosophy thesis entitled “Embryonic Development and Nutritional Requirements of Kutum Fry, Rutilus frisii kutum” in accordance with Universiti Pertanian Malaysia (Higher Degree) Act 1980 and Universiti Pertanian Malaysia (Higher Degree) Regulations 1981. The Committee recommends that the candidate be awarded the relevant degree. Members of the Examination Committee are as follows: Mihdzar Bin Abdul Kadir, PhD Associate Professor Faculty of Agriculture Universiti Putra Malaysia (Chairman) Mustafa Kamal Abdul Satar, PhD Lecture Faculty of Agriculture Universiti Putra Malaysia (Internal Examiner) Annie Christianus, PhD Lecture Faculty of Agriculture Universiti Putra Malaysia (Internal Examiner) Mohd Azmi Ambak, PhD Professor Institute of Tropical Aquaculture College Universiti of Science and Technology Malaysia (Kustem) (External Examiner) HASANAH MOHD GHAZALI, PhD Professor / Deputy Dean School of Graduate Studies Universiti Putra Malaysia Date:

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This thesis submitted to the Senate of Universiti Putra Malaysia and has been accepted as fulfilment of the requirement for the degree of Doctor of Philosophy. The members of the Supervisory Committee are as follows:

Che Roos Saad, PhD Associate Professor Faculty of Agriculture Universiti Putra Malaysia (Chairman) Abdul Razak Alimon, PhD Professor Faculty of Agriculture Universiti Putra Malaysia (Member) Ali Nikkhah, PhD Professor Faculty of Agriculture Universiti Tahran (Member) AINI IDERIS, PhD Professor / Dean School of Graduate Studies Universiti Putra Malaysia

Date:

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DECLARATION

I hereby declare that the thesis is based on my original work except for quotations and citations which have been duly acknowledged. I also declare that it has not been previously or concurrently submitted for any other degree at UPM or other institutions.

DAVOUD TALEBI HAGHIGHI

Date:

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TABLE OF CONTENTS Page DEDICATION іі ABSTRACT ііі ABSTRAK vii ACKNOWLEDGEMENTS xii APPROVAL xiv DECLARATION xvі LIST OF TABLES xx LIST OF FIGURES xxii LIST OF ABBREVIATIONS xxiv CHAPTER I GENERAL INTRODUCTION 1 Statement of Problems 4 The Significance of Study 5 Objectives 8 II LITERATURE REWIEW 9 Common Names 9 Systematic 10 Morphomerstic characteristics 11 Size 12 Color 12 Sexual dimorphism 13 Distribution 13 Fishing and Restocking 13 Threatener Factors 15 Ecological Importance 16 Digestive System 17 Larval Feeding 18 Feeding Time 20 Importance of Protein 21 Importance of Lipid 23 III GENERAL MATERIALS METHODS 26 The Proximate Analysis 26 Crude Protein 27 Crude Lipid 27 Moisture 28

xviii

Ash 29 Crude Fiber 29 Nitrogen-Free Extract 30 Gross Energy 30 Feed Preparation 32 Linear Program 33 Water quality 34 Statistical Analysis 35 IV PRELIMINARY STUDIES ON ARTIFICIAL REPRODUCTION AND SOME BIOLOGICAL CHARACTERISTICS OF KUTUM (Rutilus frisii kutum) 37 Introduction 37 Materials and Methods 41 Results 46 Length 47 Weight 47 Age 48 Fecundity 50 Discussion 52 Conclusion 57 V EMBRYONIC DEVELOPMENT OF KUTUM (Rutilu frisii kutum) 59 Introduction 59 Materials and Methods 61 Results 64 Fertilization 69 Cleavage 70 Blastula 73 Gastrula 75 Segmentation 78 Organogenesis 81 Hatching 82 VI FEEDING REGIME SPECTRUM OF KUTUM (Rutilus frisii kutum) 85 Introduction 85 Materials and Methods 89 Experimental Fish 89 Ponds Preparation 89 Experimental Diets 90 Stomach Contents Analysis 91 Results 93 Discussion 103

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VII EFFECTS OF DIFFERENT LEVELS OF CRUDE PROTEIN IN PRACTICAL DIETS ON GROWTH OF KUTUM FRY (Rutilus frisii kutum) 110 Introduction 110 Materials and Methods 112 Experimental Fish 112 Clture System 112 Diets and Feeding 113 Diets Preparation 114 Chemical Analysis 115 Statistical Analysis 115 Results 116 Discussion 123 Conclusions 132 VIII EFFECTS OF DIFFERENT LEVELS OF CRUDE LIPID IN PRACTICAL DIETS ON GROWTH OF KUTUM FRY (Rutilus frisii kutum) 133 Introduction 133 Materials and Methods 136 Experimental Fish 136 Experimental Design 137 Experimental Diets 138 The Whole body Analysis 139 Statistical Analysis 140 Results 141 Discussion 150 Conclusion 159 IX GENERAL DISCUSSION, CONCLUSION AND RECOMMENDATIONS 160 REFFRENCES 174 BIODATA OF THE AUTHOR 198

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LIST OF TABLES

Table Page 4.1 Relationships between age, length and weight with 46

absolute and relative fecundities of kutum (Rutilus frisii kutum) 4.2 Comparative of length of kutum during in recent 50 years 53 5.1 Summery of different stages of embryonic development 64

kutum fish (Rutilus frisii kutum) 5.2 Variations of egg weight, diameter and occurrence time for 64

embryonic development of kutum 6.1 Mean weight and length of kutum fry in rearing period of ponds 93 6.2 Growth performance of the kutum fry fed artificial and natural 95

food under earthen pond culture in 84 days 6.3 Zooplankton ُcommunity in gut contents of kutum fry 97 6.4 Phytoplankton community in gut contents of kutum fry 97 7.1 Ingredients and nutrients composition of the experimental 114

diets 7.2 Growth and feed conversion of kutum fry fed diets 116

containing different protein levels in 71 days 7.3 Biochemical composition of kutum larvae fed different 122

dietary protein Levels 8.1 Ingredients and proximate composition (g/100 g dry weight) 139 of experimental diets 8.2 Growth performance of kutum fry fed diets 142 containing different Lipid levels with iso-nitrogenousover 11 weeks at 22-24 °C 8.3 Morphological measurements of kutum fry fed diets 147

containing different lipid levels for 81 days

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8.4 Proximate analysis of the whole body of kutum fry fed 148 the diets containing different lipid levels (% of wet weight basis) .

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LIST OF FIGURES

Figures Page 2.1 Kutum (Rutilus frisii kutum) 9 4.1 Relationships between absolutes and relative fecundity 47

with different lengths of kutum 4.2 Relationships between absolutes and relative fecundity 48

with different weights of kutum 4.3 Relationships between absolutes and relative fecundities 50

with different ages of kutum 4.4 Relationships between percentages of ovary weight with 51

different age and weight of kutum 5.1 Mean weight, diameter and time of different stages of 65

kutum embryo development. 5.2 Stages of embryonic development of the kutum 66 5.3 Stages of morphogenesis of the kutum 67 5.4 Histology of stages embryonic development of the kutum 68 5.5 Variations of weight and diameter of kutum egg at cleavage 71

stage 5.6 Variations of weight and diameter of kutum egg at blastula 73

stage 5.7 Variations of weight and diameter of kutum egg at gasrula 75

stage 5.8 Variations of weight and diameter of egg at Segmentation 79

stage 5.9 Variations of weight and diameter of egg at organogenesis 81

stage 5.10 Variations of weight and diameter of kutum egg at hatching 83

stage 6.1 Total weight and length of Kutum fry during rearing period 94

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at pond 1 6.2 Total weight and length of Kutum fry during rearing period 94

at pond 2 6.3 Feeding items of spectrum of kutum fry in first rearing period 98 6.4 Feeding items of spectrum of kutum fry in second rearing period 99 6.5 Feeding items of spectrum of kutum fry in third rearing period 100 6.6 Feeding items of spectrum of kutum fry in fourth rearing period 101 6.7 Feeding items of spectrum of kutum fry in fifth rearing period 102 7.1 The effect of dietary protein levels on weight gain of kutum fry 118 7.2 Regression relationships between different protein levels and 119

specific growth rate (SGR) of kutum fry 7.3 Regression relationships between different protein levels and 120

feed conversion ratio (FCR) of kutum fry 7.4 The effect of different protein levels (%) on protein 121

conversion efficiency (PCE) and energy conversion efficiency (ECE)

8.1 Regression relationships between dietary of lipid levels with 144

weight gain (WG) of kutum fry 8.2 Regression relationships between dietary of lipid levels with 144

specific growth rate (SGR) of kutum fry 8.3 Regression relationships between dietary of lipid levels with 145

of specific growth rate kutum 8.4 Regression relationships between dietary of lipid levels with 145

feed conversion rate (FCR) of kutum fry

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LIST OF ABBREVIATIONS A Anal

AF Absolute Fecundity

ANOVA Analysis of Variance

AP Anterior- Posterior

Cm centimeter

CF Condition Factor

CL Crude Lipid

CP Crude Protein

CRD Complete Randomized Design

D Dorsal

DEL Deep Cell Layer

DMRT Duncan Multiple Range Test

DMSO Dimethyl Sulfoxide

DO Dissolved Oxygen

DV Dorsal- Ventral

DW Dry Weight

Epine Epinephalus

ECE Energy Conversion Efficiency

EPL Early Postlarvae

EFA Essential Fatty Acid

FCR Feed Conversion Ratio

G gram

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GnRh Gonadotropin hormone

H Height

H hours

HUFA Highly Unsaturated Fatty Acids

Kg Kilogram

K cal Kilocalorie

L. L Late larvae

L Liter

L Length

L3 Cubic Length

LINDO Linear Interactive and Discrete Optimizer

LP Linear Programming

M meter

m2 Square meter

mm2 Square millimeter

m3 cubic meter

MBW Mean Body Weight

MD Mean Development

Mg milligram

Min minutes

Mm millimeter

Ml milliliter

Μm micronmeter

MUFA Monounsaturated fatty acids

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NaOH Sodium hydroxide

NRC National Research Council

PER Protein Efficiency Ratio

PCE Protein Conversion Efficiency

PL Postlarval

P/L Protein/ Lipid ratio

PS Protein Sparing

PUFA Polyunsaturated Fatty Acids

RF Relative Fecundity

R regression

SD Standard Deviation

SE Standard Error

SFA Saturated Fatty Acids

SGR Specific Growth Rate

SR Survival Rate

W Weight

WG Weight Gain

YS Yolk Sac

YSL Yolk Syncytial Layer