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EMBRYONIC DEVELOPMENT AND NUTRITIONAL REQUIREMENTS OF KUTUM FRY, RUTILUS FRISII KUTUM
DAVOUD TALEBI HAGHIGHI
DOCTOR OF PHILOSOPHY UNIVERSITI PUTRA MALAYSIA
2006
i
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
ii
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
iii
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
iv
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
v
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.
vi
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.
vii
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
viii
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)
ix
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
x
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%.
xi
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%.
xii
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.
xiii
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.
xiv
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:
xv
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:
xvi
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:
xvii
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
xix
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
xx
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
xxi
8.4 Proximate analysis of the whole body of kutum fry fed 148 the diets containing different lipid levels (% of wet weight basis) .
xxii
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
xxiii
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
xxiv
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
xxv
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
xxvi
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