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Christian E. W. Steinberg
Aquatic
Animal
Nutrition
A Mechanistic Perspective from
Individuals to Generations
Aquatic Animal Nutrition
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Aquatic Animal Nutrition

A Mechanistic Perspective from Individuals
to Generations
Department of Biology
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Humboldt University at Berlin

Berlin, Germany
ISBN 978-3-319-91766-5 ISBN 978-3-319-91767-2 (eBook)

https://doi.org/10.1007/978-3-319-91767-2
Library of Congress Control Number: 2018953153
© Springer Nature Switzerland AG 2018

This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of
the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation,
broadcasting, reproduction on microfilms or in any other physical way, and transmission or information
storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology
now known or hereafter developed.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication
does not imply, even in the absence of a specific statement, that such names are exempt from the relevant
protective laws and regulations and therefore free for general use.
The publisher, the authors and the editors are safe to assume that the advice and information in this book
are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the
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This Springer imprint is published by the registered company Springer Nature Switzerland AG
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
Preface
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Never attend an expedition to Vietnam that is devoted to the study of freshwater and
marine fishes! However, if you do, you run the risk of making good new friends in
the commercial and ornamental fish business, sharing some (too many!) Tiger beers
and promising to spread new information about the nutrition of fish and other
aquatic animals. I took that risk, went to Vietnam, made new friends, and fell in love
with South (East) Asia. Several years have passed since then, and experts, as well as
laypersons, have had to tolerate and survive several seminars on aquatic animal
nutrition, given by me. Now, it is time to keep my promise.
A freshwater ecologist by education and a stress ecologist by preference, my
primary interest has not been to write a book that discusses higher productivity in
the aquaculture industries or reviews recipes for more effective functional aquafeeds
to increase survival, reproduction or productivity of farmed animals. Instead, I am
more interested in answering the question of how certain dietary ingredients influ-
ence the life history traits not only of the consumers but also of their succeeding
generations. In evolutionary-ecological terms: How do dietary components impact
the Darwinian fitness of populations and thereby influence their long-term persis-
tence in the ecosystem? The ecologist in me always noticed gaps in the more prag-
matic experimental approaches of raising aquatic animals referenced in this book.
To identify the gaps, I had to sometimes crawl ashore, since I felt obliged to borrow
information about new developments from terrestrial or laboratory model animal
studies. Nevertheless, I hope that my raised forefinger will encourage the develop-
ment of new experimental setups for the aquaculture community.
The content for the originally planned one-volume book on “Aquatic Animal
Nutrition,” however, turned out to be so voluminous that I split it into two volumes.
I thank Springer Publishing Company for this courtesy. My sincere appreciation
goes particularly to “my Springer ladies” in Dordrecht, namely Alexandrine
Cheronet and Judith Terpos, who were always very supportive and never hesitated
to answer my questions, even if they were simple.
Furthermore, I am thankful to all the photographers and artists who allowed me
to use their wonderful images free of charge. Doubtless, they contributed to an
attractive appearance of this book. We all agree that good illustrations can often
v
vi Preface
explain complex ideas much better than thousands of words. Nevertheless, also
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good words count, and I thank Sarah L. Poynton for the excellent word crafting that
resulted in the title.
Even to a book, space limitation applies. Due to this circumstance, I would like
to apologize in advance to all individuals whose research was not cited or whose
papers have not been discussed in full but whose work has certainly advanced the
understanding of this complex field of research, practice, and education.
This book is dedicated to my bright grandkids, Anna S. and Paul N., who like
watching colorful and intriguing fishes in a living room tank. Since they started this
business at a much younger age than I did, I am certain that one, or both of them,
will be inclined to write brilliant books on this fascinating subject.
Berlin, Germany Christian E. W. Steinberg

Contents
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1 Introduction – ‘You Are What You Eat’................................................... 1

Appendix.................................................................................................... 6
Technical Note....................................................................................... 6
References.................................................................................................. 7
2 Diets and Digestive Tracts – ‘Your Food Determines
Your Intestine’........................................................................................... 9
2.1 Digestive Tract.................................................................................. 9
2.2 Digestion........................................................................................... 14
2.2.1 Protein Digestion.................................................................. 15
2.2.2 Lipid Digestion..................................................................... 16
2.2.3 Carbohydrate Digestion........................................................ 16
2.3 Ontogenesis and the Intestine........................................................... 17
2.3.1 Fishes.................................................................................... 17
2.3.2 Invertebrates.......................................................................... 26
2.4 Herbivory, a Disadvantageous Acquization Strategy?...................... 34
2.4.1 Fishes.................................................................................... 37
2.4.2 Invertebrates.......................................................................... 40
2.5 Starvation and Gut Morphology....................................................... 42
2.6 Trophic Positions: An Omnivores’ Dilemma?.................................. 45
2.7 Concluding Remarks......................................................................... 53
References.................................................................................................. 54
3 The Intestinal Microbiota – ‘Your Eating Feeds a Plethora
of Guests’ and ‘This Plethora of Guests Determines
Who You Are and How Well You Do’........................................................ 61
3.1 Invertebrates...................................................................................... 67
3.1.1 Hydrozoa............................................................................... 67
3.1.2 Mollusks................................................................................ 68
3.1.3 Echinoderms......................................................................... 72
3.1.4 Crustaceans........................................................................... 73
vii
viii Contents
3.2 Fishes................................................................................................ 80

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3.2.1 Microbiome Ontogenesis...................................................... 81

3.2.2 Does a Core Microbiome Exist?........................................... 85
3.2.3 Zebrafish as Witness of Microbiome Development.............. 111
3.2.4 Control Functions by Gut Microbiota................................... 115
References.................................................................................................. 128
4 Dietary Restriction, Starvation, Compensatory
Growth – ‘Short-Term Fasting Does Not Kill You:
It Can Make You Stronger’....................................................................... 137
4.1 Indicators of Starvation..................................................................... 143
4.2 Starvation Tolerance and Starvation Impact..................................... 148
4.2.1 Cnidarians............................................................................. 162
4.2.2 Rotifers.................................................................................. 163
4.2.3 Mollusks................................................................................ 166
4.2.4 Echinoderms......................................................................... 168
4.2.5 Crustaceans........................................................................... 170
4.2.6 Fishes.................................................................................... 192
4.2.7 Summary of Starvation Effects............................................. 198
4.2.8 Starvation: Point-of-no-Return............................................. 199
4.3 Compensatory Growth...................................................................... 201
4.3.1 Invertebrates.......................................................................... 206
4.3.2 Fishes.................................................................................... 214
4.4 Compensatory Growth in Populations.............................................. 228
4.5 Regulation of Compensatory Growth............................................... 230
4.5.1 Appetite-Regulating Hormones............................................ 233
4.5.2 Neuropeptides....................................................................... 236
4.5.3 Transcription of Growth Regulators..................................... 243
4.6 Concluding remarks.......................................................................... 252
References.................................................................................................. 255
5 Chrononutrition – ‘The Clock Makes Good Food’................................ 289
5.1 How Does a Biological Clock Work?............................................... 293
5.1.1 Fishes.................................................................................... 294
5.1.2 Invertebrates.......................................................................... 302
5.2 Food and Circadian Gene Transcription........................................... 308
5.2.1 Major Nutrients..................................................................... 308
5.2.2 Xenobiotic or Antinutritional Compounds........................... 315
References.................................................................................................. 325
Contents ix
6 Transgenerational Effects – ‘Your Offspring Will Become

What You Eat’............................................................................................ 333
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6.1 Parental Effects................................................................................. 335

6.1.1 Maternal Effects.................................................................... 344
6.1.2 Paternal Effects..................................................................... 373
6.2 What Is Epigenetics?......................................................................... 378
6.2.1 Time Scales of Epigenetic Inheritance.................................. 380
6.2.2 Epigenetic Mechanisms........................................................ 381
References.................................................................................................. 416
7 Trophic Diversification and Speciation – ‘Your Eating
Fuels Evolution’........................................................................................ 431
7.1 Individual Specialization.................................................................. 437
7.2 Underlying Mechanisms of Speciation............................................. 438
7.3 Trophic Speciation............................................................................ 441
7.3.1 Ancient Lakes....................................................................... 441
7.3.2 Rivers.................................................................................... 451
7.3.3 Coral Reefs............................................................................ 452
7.4 Convergent Evolution....................................................................... 454
7.5 Rapid Speciating Taxa...................................................................... 458
7.5.1 Pumpkinseed Sunfish............................................................ 458
7.5.2 Cyprinodon........................................................................... 459
7.5.3 Terapontidae (Grunters)........................................................ 460
7.5.4 Three-Spined Stickleback..................................................... 460
7.5.5 Arctic Charr.......................................................................... 462
7.6 Time Span of Trophic Speciation..................................................... 465
References.................................................................................................. 467
Abbreviations and Glossary
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AAs amino acids

abcb4 gene encoding multidrug resistance protein 3, a membrane-
bound transporter
abcg2 gene encoding ATP-binding cassette subfamily G member
2, a membrane-bound tranporter
Acrophase time at which the peak of a rhythm occurs
AgRPs agouti-related proteins: neuropeptides produced in the
brain with an appetite-stimulating potential; activated by
the hormone ghrelin, inhibited by the hormone leptin
AHR anti-hydroxyl radical
AhR aryl-hydrocarbon receptor, Ah receptor: a ligand-activated
transcription factor involved in the regulation of biological
responses to planar aromatic (aryl) hydrocarbons, includ-
ing xenobiotic compounds
ahr2 gene encoding the aryl-hydrocarbon receptor 2 found in
zebrafish
AL ad libitum = “at one’s pleasure”
Alternative splicing a regulated process during gene expression that results in a
single gene coding for multiple proteins
ambra1a, ambra1b autophagy-regulating genes
AMPK AMP-activated protein kinase: 5’adenosine
monophosphate-activated protein kinase, an enzyme cen-
tral in cellular energy homeostasis
anadromous fish migrate from the sea up into freshwater to spawn;
examples are salmon and striped bass
anorexigen appetite-suppressing drug or food constituent
apaf-1 encodes apoptotic protease-activating factor 1, a pro-
aptotic protein
arα, arβ encode androgen receptor α and β
ASA anti-superoxide anion
xi
xii Abbreviations and Glossary
Autocrine (signaling) a form of cell signaling in which a cell secretes a signaling

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chemical that binds to receptors on that same cell

Autophagy natural, regulated, destructive mechanism of the cell that
disassembles unnecessary or dysfunctional components
BAX Bcl-2-associated X protein: functions as part of an apop-
totic activator
BBS bombesin-like peptides: a large family of peptides initially
isolated from amphibian skin; peptides with neuroendo-
crine and neuromodulator function in fish
Bcl2 B-cell leukemia/lymphoma-2: protein regulating cell death
(apoptosis), by either inducing (pro-apoptotic) or inhibit-
ing (anti-apoptotic) apoptosis
bd β-defensins: antimicrobial peptides implicated in the resis-
tance of epithelial surfaces to microbial colonization
beclin autophagy-regulating gene
Bioaccumulation the process which causes chemical concentrations in the
tissues of an aquatic organism to exceed those in the water,
due to uptake by all exposure routes
bmal(1,2..) encode transcription factor(s) involved in the circadian
rhythm (clock gene); together with clock the positive loop
of the molecular clock
bmp15 encodes bone morphogenetic factor 15, mainly involved in
folliculogenesis
C3 complement component 3, a protein of the immune sys-
tem, central in the activation of the complement system; a
connecting link between innate and acquired immunity
CART cocaine and amphetamine regulated transcript: a neuro-
peptide that produces similar behavior in animals to
cocaine and amphetamine having roles in reward, feeding,
and stress
caspase-3…9 members of the cysteine-aspartic acid protease (caspase)
family: sequential activation of caspases plays a central
role in the execution-phase of cell apoptosis
CAT catalase: enzyme that catalyzes the dismutation of hydro-
gen peroxide (H2O2) to water (H2O) and oxygen (O2)
Catch-up growth attainment of control size
CCK cholecystokinin: peptide hormone of the gastrointestinal
system stimulating the digestion of fat and protein
cenpf1...3 encode centromere proteins F, involved in physical cell
division
c-fos FBJ murine osteosarcoma viral oncogene homolog, a
proto-oncogene involved in signal transduction, cell prolif-
eration and differentiation
CFU colony-forming units
CG compensatory growth: faster than usual growth rate
Abbreviations and Glossary xiii
Chaperon protein stabilizing new proteins to ensure correct folding

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or helping to refold proteins that were damaged by cell

stress
chordin early developmental gene ruling the dorsalizing process
CK creatine kinase: enzyme that catalyzes the conversion of
creatine and utilizes adenosine triphosphate (ATP) to cre-
ate phosphocreatine (PCr) and adenosine diphosphate
(ADP)
clk, clock encodes transcription factor(s) involved in the circadian
rhythm (clock gene); together with bmal(1,2..) the positive
loop of the molecular clock
Complement factor B an acute-phase protein increasing during inflammation
Cosinor analysis analysis of biologic time series that demonstrate pre-
dictible rhythms
cox-2 encodes cyclooxygenase forming prostanoids from PUFAs
CRH corticotropin-releasing hormone
cry encodes cryptochrome, blue light-sensitive flavoproteins
involved in the circadian rhythm; with per2 and tim the
negative loop of the molecular clock
CSF-1R colony-stimulating factor 1 receptor (also macrophage
colony-stimulating factor receptor): receptor for a cytokine
called colony-stimulating factor 1, which controls the pro-
duction, differentiation, and function of macrophages
CTR calcitriol, increasing the uptake of calcium from the gut
into the blood
Curcumin complex polyphenol in the rhizome of turmeric, Curcuma
longa
CuZn-SOD copper/zinc superoxide dismutase; enzyme that catalyzes
the dismutation of superoxide radical (•O2−) to water (H2O)
and oxygen (O2)
cyc encodes cycle protein; genetic transcription-translation
feedback loop that generates circadian rhythms
cyp17-II cytochrome P450 c17II; a monooxygenase affecting
growth, gonad differentiation and development, and other
reproductive traits of fish
cyp1a gene encoding cytochrome P450, family 1, subfamily A, is
involved in phase I xenobiotic and drug metabolism
Cytokine cell signaling protein
DAF-16 ortholog of the FOXO family of transcription factors in the
nematode C. elegans; it is the primary (but not the only)
transcription factor required for lifespan extension
dazl deleted in azoospermia-like: This gene encodes a member
of the depleted in azoospermia-like (DAZL) protein
family
xiv Abbreviations and Glossary
dbt encodes DOUBLETIME protein; a kinase that phosphory-

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lates PER protein that regulates the molecularly driven,

biological clock controlling circadian rhythm
DE digestible energy
DGGE denaturing gradient gel electrophoresis; method for identi-
fying genes from natural ecosystems
Dicer a key initiative protein of the RNA interference (RNAi)
pathway: (= endoribonuclease dicer) an enzyme cleaves
double-stranded RNA (dsRNA) and pre-microRNA (pre-
miRNA) into short double-stranded RNA fragments called
small interfering RNA
DM dry matter
DNMT DNA methyltransferase
dpf days past fertilization
dph days past hatch
DR dietary restriction
dsRNA double-stranded (ds)RNA, central in RNAi
Dysbiosis microbial imbalance or maladaptation
EF-1α eukaryotic translation elongation factor 1 alpha responsi-
ble for the enzymatic delivery of aminoacyl tRNAs to the
ribosome
EFA essential fatty acid; EFA requirements vary qualitatively as
well as quantitatively among different animal species
EGCG epigallocatechin-3-gallate, the most active ingredient in
green tea
ELOVL1 FA elongase elongates saturated and monounsaturated C20-
C26 acyl-CoAs
ELOVL2 FA elongase elongates C20-C22 polyunsaturated acyl-CoAs
ELOVL3 FA elongase elongates saturated and unsaturated C16-C22
acyl-CoAs
ELOVL4 FA elongase for the synthesis of ULCFAs (C ≥ 26)
ELOVL5 FA elongase elongates C18-C20 polyunsaturated acyl-CoAs
ELOVL6 FA elongase elongates C12:0-C16:0
ELOVL7 FA elongase elongates saturated and unsaturated C16-C22
acyl-CoAs
endozoochory dispersal of spores or seeds in the gut of animals
Enterocytes intestinal absorptive cells: simple columnar epithelial cells
found in the small intestine
EPA eicosapentaenoic acid, an omega-3 fatty acid with the
chemical formula C20H30O2
ER estrogen receptor
FA fatty acid
FAA food anticipatory activity
fas encodes fatty acid synthase
Abbreviations and Glossary xv
fMHC fast myosin heavy chain, ATP-dependent motor protein,

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involved in muscle contraction and other motility pro-

cesses in eukaryotes
FoxOs forkhead box proteins, a family of transcription factors
regulating the expression of genes involved in cell growth,
proliferation, differentiation, and longevity
fsh encodes follicle-stimulating hormone
fshr encodes follicle-stimulating hormone receptor
GABA γ-aminobutyric acid, a non-proteinogenic amino acid:
inhibitory neurotransmitter
GAL (Gal) galanin: neuropeptide involved in feeding and growth
Garcinol polyisoprenylated benzophenone derivative isolated from
kokum, Garcinia indica
gdh (also gldh) encodes glutamate dehydrogenase
Geldanamycin antitumor antibiotic inhibiting the function of HSP90
Genistein isoflavonoid phytoestrogen
germ-free animals animals that have no microorganisms living in or on them
GF germ free, gnotobiotic
GH growth hormone that activates AgRP, thus increasing appe-
tite; is opposed by the hormone leptin; furthermore, as
pleotropic hormone GH is involved in growth, stress
response, energy homeostasis, reproduction
GHR growth hormone receptor; a protein that is a transmem-
brane receptor for growth hormone
GHra, GHrb growth hormone receptors a & b
GHRH growth hormone-releasing hormone: a releasing hormone
of growth hormone (GH) that stimulates GH production
and release by binding to the GHRH Receptor (GHRHR)
on cells in the anterior pituitary
Ghrl also ghrelin: growth hormone release inducing: the “hun-
ger hormone” is a peptide hormone regulating appetite and
the distribution and rate of use of energy. Ghrelin acts as
orexigenic hormone
GHS-R growth hormone secretagogue receptor, or ghrelin recep-
tor, is a ghrelin-binding receptor and plays a role in energy
homeostasis and regulation of body weight
GI gastrointestinal (tract)
GIFT genetically improved farmed tilapia by conventional
breeding
GLP glucagon-like peptide, a neuropeptide
GnRHs gonadotropin-releasing hormones; among other functions:
decreases quantity and frequency of food consumption
GO gene ontology
goosecoid early developmental gene ruling the dorsalizing process
GPx glutathione peroxidase
xvi Abbreviations and Glossary
GR glutathione reductase
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GRP gastrin-releasing peptide: a neuropeptide, stimulating the

release of gastrin from the G cells of the stomach
GST glutathione transferase
gstr1 gene encoding glutathione transferase rho1
HCRT hypocretin (orexin) neuropeptide precursor regulating
appetite
Heterochrony developmental change in the timing or rate of events, lead-
ing to changes in size and shape
hpf hours past fertilization
HSFs heat shock factors: transcription factors regulating the
expression of heat shock proteins
hsl hormone-sensitive lipase gene
HSP90 heat shock protein 90 (atomic mass approximately 90 kDa),
a chaperone protein assisting other proteins to fold prop-
erly, stabilizing proteins against stress; it also canalizes
phenotypic variability
HSPs heat shock proteins: family of proteins produced by cells in
response to exposure to stressful conditions. Many HSPs
perform chaperone function
HYP hypothalamus
IGF-1…3 insulin-like growth factor 1…3, hormones similar in
molecular structure to insulin
IgM immunoglobulin M
InAP intestinal alkaline phosphatase
irf7 encodes interferon regulatory factor 7, a transcription fac-
tor, central in the transcriptional activation of virus-
inducible cellular genes, including the type I interferon
genes
Isothiocyanate chemical group –N=C=S; mustard oils contain
isocyanates
IU international units: amount of a drug, hormone, vitamin,
enzyme, etc., that produces a specific effect as defined by
an international body and accepted internationally
KEGG Kyoto Encyclopedia of Genes and Genomes, a database
resource for understanding high-level functions and utili-
ties of the biological system
kiss1, kiss2 encode kisspeptins
Kisspeptins peptides that stimulate gonadotropin release
lc3 autophagy-related gene
LDLR low-density lipoprotein receptor, mediates the endocytosis
of cholesterol-rich LDL and thus maintains the plasma
level of LDL
Abbreviations and Glossary xvii
Lecithotrophy nourishment and development of the embryo only via the

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yolk originally contained within its egg; opposite:

matrotrophy
LEP leptin, a hormone that regulates appetite to achieve energy
homeostasis by inhibiting hunger; it is opposed by the hor-
mone ghrelin
lh gene encoding the luteinizing hormone
lhcgr gene encoding the luteinizing hormone/choriogonadotro-
pin receptor
Lipofuscin finely granular yellow-brown pigment granule indicative
of aging cells
lpl encodes lipoprotein lipase
LPS lipopolysaccharides (also lipoglycans, endotoxins): large
molecules consisting of a lipid and a polysaccharide; found
in Gram-negative bacteria and cyanobacteria and elicit
strong immune responses in animals
Lysozyme c glycoside hydrolase, an enzyme that damages bacterial cell
walls
Matrotrophy form of maternal care during organism development, in
which the embryo is supplied with additional nutrition
from the mother, e.g. through a placenta; opposite:
lecithotrophy
MC4R melanocortin-4 receptor, a protein involved in feeding
behavior, suppressing hunger
mch encodes melanin-concentrating hormone: a cyclic orexi-
genic hypothalamic peptide originally isolated from the
pituitary gland of teleost fish
Melatonin N-acetyl-5-methoxy tryptamine, a hormone that antici-
pates the daily onset of darkness; hormone affecting the
modulation of wake/sleep patterns
mgst3a gene encoding microsomal glutathione transferase 3a (in
zebrafish), central in phase II detoxification
mhc2a (also mhcIIa) encodes mhc class 2A chain (major histocompability com-
plex IIα), molecules found on antigen-presenting cells,
important in initiating immune responses
Microbiota resident microbial communities in fishes and invertebrates
microRNA miRNA, about 22 nucleotides, functioning in RNA silenc-
ing and posttranscriptional regulation of gene expression
Mn-SOD manganese superoxide dismutase; an enzyme that cata-
lyzes the dismutation of superoxide radical (•O2−) to water
(H2O) and oxygen (O2)
MyoD one myogenic regulatory factor
myog encodes myogenin
xviii Abbreviations and Glossary
Myogenin (= myog, myogenic factor 4) transcription factor involved

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in the coordination of skeletal muscle development or

myogenesis and repair
Myostatin a myokine: protein produced and released by myocytes
inhibiting muscle cell growth and differentiation, member
of the TGF-β superfamily
myp encodes major yolk protein
NAD nicotinamide adenine dinucleotide
Neurolipofuscin lipofuscin in the nervous system
NF-κB nuclear transcription factor-κB: protein complex that con-
trols transcription of DNA, cytokine production and cell
survival; found in almost all animal cell types and involved
in cellular responses to stimuli such as stress, cytokines,
free radicals, ultraviolet irradiation, oxidized low-density
lipoprotein, and bacterial or viral antigens
NPY neuropeptide Y: with slight variations, a neurotransmitter
in the brain and in the autonomic nervous system of ani-
mals; one of the strongest orexigenic signals
opn encodes osteopontin, an important mediator of bone (re)
modeling
Orexigen drug, hormone, or compound that stimulates appetite
osr-1/unc-43/sek-1 pathway that promotes resistance to osmotic stress; osr-1
is coupled to SEK-1 (a MAK kinase) through UNC-43
(Ca2+/calmodulin-dependent protein kinase II)
osx encodes osterix, a transcription factor for osteoblast dif-
ferentiation, mediates antitumor activity in murine
osteosarcoma
OTU operational taxonomic units
OX orexin, a neuropeptide that regulates arousal, wakefulness,
and appetite
Oxidative burst refer to respiratory burst
Oxidative stress states where the balance between generation and elimina-
tion of ROS is disturbed in favor of the generation of ROS
p38 encodes P38 mitogen-activated protein kinase; is activated
by a variety of cellular stresses
paqr8 encodes progestin and adipoQ receptor family
Paracrine (signaling) a form of cell-cell communication in which a cell produces
a signal to induce changes in nearby cells; opposite:
autocrine
PBS phosphate-buffered saline (buffer)
PC phosphatidylcholine
pepT1…2 encode oligopeptide transporters (members of the solute
carrier family 15) encode solute carriers localized to the
brush border membrane of the intestinal epithelium and
mediate the uptake of di- and tripeptides
Abbreviations and Glossary xix
per1,2 encoding the period circadian regulators 1,2; with cry1 and
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tim the negative loop of the molecular clock

PG phosphatidylglycerol
PI phosphatidylinositol
Pineal organ pineal body, epiphysis cerebri, epiphysis or the “third eye”:
small endocrine gland producing melatonin
PIT pituitary or hypophysis, an endocrine gland. Its hormones
help control: growth, blood pressure, and certain functions
of the sex organs, thyroid glands, metabolism, and
reproduction
POA preoptic area, part of the hypothalamus; responsible for
thermoregulation and receives nervous stimulation from
thermoreceptors in the skin, mucous membranes, and
hypothalamus itself
POMC proopiomelanocortin, a precursor polypeptide which
cleavage gives rise to several peptide hormones
pparα…γ encode peroxisome proliferator-activated receptors α, β, γ;
nuclear receptor proteins functioning as transcription fac-
tors; central in the regulation of cellular differentiation,
development, and metabolism (carbohydrate, lipid,
protein)
PSM plant secondary metabolites
Procyanidin a condensed tannin
proPO prophenoloxidase: part of the major innate defense system
in invertebrates via melanization of pathogens and dam-
aged tissues
PUFA polyunsaturated fatty acid containing two or more ethyl-
enic bonds, such eicosapentaenoic acid (EPA, 20:5n─3 or
20:5ω─3)
PY peptide Y, a NPY-related peptide
PYY peptide YY is a member of the neuropeptide Y (NPY)
family
QTL quantitative trait locus: section of DNA, the locus that cor-
relates with variation in a phenotype, the quantitative trait
Quercetin a flavonoid polyphenol
Rapamycin also sirolimus; chemical immune suppressant
RAS recirculating aquaculture systems
Respiratory burst rapid production and release of reactive oxygen species
Resveratrol a stilbenoid polyphenol
retinoic acid a metabolite of vitamin A (retinol) that mediates the func-
tions of vitamin A required for growth and development
Ribotype molecular bacterial identification using information from
rRNA-based phylogenetic analyses
xx Abbreviations and Glossary
RNAi RNA interference or RNA silencing: biological process in

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which RNA molecules inhibit gene expression or transla-

tion, by neutralizing targeted mRNA molecules
rnf213 encodes E3 ubiquitin-protein ligase RNF213, involved in
protein ubiquitination
ROS reactive oxygen species: chemical reactive molecules con-
taining oxygen, such as peroxides, superoxide, hydroxyl
radical, and singlet oxygen (also see electrophilic stress
and oxidative stress)
RXR retinoid receptors: nuclear receptors that bind to retinoids;
when bound to a retinoid, they act as transcription factors
SAH S-adenosyl-l-homocysteine: amino acid derivative, inter-
mediate in the synthesis of cysteine and adenosine; SAH is
formed by the demethylation of S-adenosyl-l-methionine
(SAM)
SAM S-adenosyl-l-methionine: involved in methyl group
transfers
SCN suprachiasmatic nucleus of the hypothalamus in mammals,
a neuronal structure defining the circadian rhythms
SGR specific growth rate
SIRT1 sirtuin (silent mating type information regulation 2 homo-
log) 1, an enzyme deacetylating proteins and thereby regu-
lating reaction to stress and longevity
SMADs structurally similar proteins that are the main signal trans-
ducers for receptors of the transforming growth factor-beta
(TGF-β) superfamily
SOD superoxide dismutase: enzyme that catalyzes the dismuta-
tion of superoxide (•O2−) radicals into ordinary molecular
oxygen (O2) and hydrogen peroxide (H2O2)
SOD1 Cu-Zn superoxide dismutase
SOD2 Mn superoxide dismutase = superoxide dismutase 2
Somatomedins group of hormones that promote cell growth and division
in response to stimulation by growth hormone (GH)
Sulforaphane organosulfur compound (isothiocyanate) in cruciferous
vegetables
sult2_st2 gene encoding the cytosolic sulfotransferase 2 (in
zebrafish)
T3 thyroid hormone; more active than T4 by a factor of 3 to 5
TAG triglyceride or triacylglycerol: ester derived from glycerol
and three fatty acids
TGF-β transforming growth factor β: multifunctional cytokine.
The TGF-β superfamily includes endogenous growth-
inhibiting proteins, for instance, with anti-inflammatory
function
Abbreviations and Glossary xxi
tim a, b, h encode timeless proteins; essential in regulating the circa-

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dian rhythm; part of a transcription-translation negative

feedback loop involving the period (per) and cry genes and
their proteins
TLRs Toll-like receptors: class of proteins central in the innate
immune system that recognize structurally conserved mol-
ecules derived from microbes
toll encodes members of the Toll-like receptor class of
proteins
TOR target of rapamycin: highly conserved, nutrient-sensitive
protein kinase, a central controller of protein synthesis,
cell growth, cell proliferation, cell motility, cell survival,
autophagy, transcription, and aging
TRH thyrotropin-releasing hormone: a releasing hormone, pro-
duced by the hypothalamus that stimulates the release of
thyrotropin
Trp tryptophan: 1 of the 22 standard amino acids and an essen-
tial in diets; distinguishing structural feature is the indole
functional group
TSH thyroid-stimulating hormone: a pituitary hormone that
stimulates the thyroid gland to produce thyroxine (T4)
usp5 encodes ubiquitin carboxyl-terminal hydrolase 5, a deu-
biquitinating enzyme
Veliger planktonic larva of many sea snails and freshwater snails,
as well as most bivalve mollusks
ZT zeitgeber time, time from any external or environmental
cue on that entrains or synchronizes an organism’s biologi-
cal rhythms to the Earth’s 24-hour light/dark cycle and
12-month cycle. The time of the cue is ZT 00:00
β-diversity ratio between regional and local species diversity
β-endorphin endogenous opioid neuropeptide and peptide hormone that
is produced in certain neurons within the central nervous
system and peripheral nervous system; even protists, such
as Tetrahymena, produce this hormone
β-oxidation catabolic process by which fatty acid molecules are broken
down in the mitochondria in eukaryotes to generate acetyl-
CoA, which enters the citric acid cycle, and NADH and
FADH2, which are coenzymes used in the electron trans-
port chain
Chapter 1
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Introduction – ‘You Are What You Eat’
Abstract The trivial word ‘You are what you eat’ (YAWYE) actually applies also
to fish and aquatic invertebrates; however, not literally, but in a more hidden, subtle
manner. This introductory chapter will briefly address the content of Volume I of
Aquatic Animal Nutrition: Chapter 2 recalls basic textbook knowledge and dis-
cusses dietary impacts on morphology and functioning of the intestine. Chapter 3
focuses on the central significance of the intestinal microbiota, the forgotten ecosys-
tem. Central in Chap. 4 is the message that dietary restriction and starvation are
natural occurrences and do not necessarily kill the individuals which often respond
with compensatory growth and improved Darwinian fitness – even of their off-
spring. Chapter 5 discusses the circadian rhythmicity of digestive and biotransfor-
mation gene transcription and questions the paradigm of ‘antinutritional factors’.
Chapter 6 addresses transgenerational dietary effects including starvation resis-
tance; and the last Chapter shows that diets can be the basis for sympatric speciation
whereby not only genetical, but also epigenetical mechanisms likely apply.
‘You are what you eat!’

This trivial, hackneyed word is certainly a consensus promoting empty phrase.
However, does its frequent usage increase the plausibility of it’s meaning? Is this
phrase really true, or is it just the wishful thinking of some zealous nutritionists who
want the majority of the human population to live healthier or take up diets such as
vegetarianism or veganism? Is it more than a consensus creating, but empty phrase?
‘Are you really what you eat?’
Let’s therefore briefly switch to a bizarre aquatic creature: the black swallower
(Chiasmodon niger), which has attracted the attention of many marine biologists.
This common species of fish has a worldwide distribution in tropical and subtropi-
cal waters, at depths of 700–2745 m (Coad and Reist 2004). The black swallower
feeds on bony fishes, which it swallows whole. Incredibly, its highly distensible
© Springer Nature Switzerland AG 2018 1

C. E. W. Steinberg, Aquatic Animal Nutrition,
https://doi.org/10.1007/978-3-319-91767-2_1
2 1 Introduction – ‘You Are What You Eat’
stomach allows it to swallow prey over twice its length and 10 times its mass
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(Fig. 1.1) (Jordan 1905).

Provided that “you are what you eat” (YAWYE) is applied literally, one could
hypothesize that the black swallower, following the consumption of prey larger than
itself, should grow and grow and eventually reach the size and appearance of its
prey. Does this actually happen? If so, the black swallower could be a real-world
equivalent to the abstract conceptualization of a frog metamorphosing into a fly – a
literal interpretation of YAWYE, as cleverly depicted by American artist, Sarah
DeRemer (Fig. 1.2).
Of course, Sarah’s peculiar creature is pure fiction, and animals generally do not
metamorphose into their dietary prey. If YAWYE is really valid as opposed to being
a handy phrase for nutritionists, it must take place at a more subtle, cryptic level.
To identify the impacts of dietary sins, different diets, or healthy eating, we have
to leave the simple phenotypic level and must become acquainted with the micro-
biological, biochemical, and biomolecular levels of the consumer – levels which, as
Fig. 1.1 A black swallower containing a fish much larger than itself. (From Günther 1880, cour-
tesy of the Biodiversity Heritage Library)
Fig. 1.2 A fly-eating frog

metamorphoses into a fly,
as depicted by the artist
Sarah DeRemer (©Sarah
DeRemer)
1 Introduction – ‘You Are What You Eat’ 3
we shall see, are not only affected by present diet, but the diets of past generations,
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even spanning through evolutionary time.

We shall see that YAWYE is certainly applicable at these levels, with complex
effects that have far reaching impacts, surpassing the individual consumer and span-
ning across generations. In this context, we can refine this trivial phrase, and instead
claim:
• ‘Your food determines your intestine’
We must briefly recall textbook knowledge about the tools by which aquatic
animals can make use of their diets. The intestine of herbivores, omnivores, and
carnivores differ significantly. Here we learn that aquatic animals – fish in
particular – are very flexible; if necessary, fish can change the morphology of the
intestine with a change in trophic niche;
• ‘Your eating feeds a plethora of guests’
A diet does not only feed the individual consumer by supplying energy, macro-,
and micronutrients, but also feeds an extremely high number of diverse microorgan-
isms in the intestine. Most of these microorganisms provide additional metabolic
pathways and, thus, utilize dietary intake more efficiently than microbe-free ani-
mals ever could. These microorganism communities can be considered as an addi-
tional and effecacious organ;
• ‘This plethora of guests determines who you are and how well you do’
By several mechanisms, beneficial microorganisms suppress adverse bacteria,
cyanobacteria, and yeasts and in addition, provide signal molecules that strengthen
immunity as well as resistance to pathogens and parasites. More surprisingly, the
composition of the intestinal microbiotia has the potential to even determine gen-
der – at least in certain invertebrates;
• ‘Short-term fasting does not kill you – it can make you stronger’
In natural habitats, food is not always available in sufficient quantities or quali-
ties. Dietary restriction (food shortage) and starvation are regular occurrences for
many animals in a variety of ecosystems. They have had to develop coping mecha-
nisms for these situations – with surprising results. For instance, parental short-term
starvation can increase disease resistance in offspring and can even have beneficial
impacts on population growth;
• ‘The clock makes good food’
From studies of terrestrial invertebrates and mammals, there is accumulating evi-
dence that biotransformation activity on the biomolecular and biochemical levels is
subject to circadian rhythmicity. Depending on the time of day, adverse effects from
identical exposure to natural or synthetic xenobiotic compounds can differ signifi-
cantly. Since this rhythmicity appears to be evolutionarily conserved, it can be
hypothesized that it will also apply to aquatic animals. The confirmation of this
hypothesis will surely challenge the paradigm of the so-called anti-nutritional

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factors in animal feed in general, and for aquafeed in particular;

• ‘Your offspring will become what you eat’
Recent studies, most of which concerning terrestrial vertebrates and inverte-
brates, have discovered transgenerational mechanisms of how diet-mediated prop-
erties of the parental generation are passed not only to the next filial generation, but
also to the grand and great grand generations. These phenomena have been detected
in terrestrial invertebrates as well as in mammals, and appear to be conserved
throughout evolutionary history; there is no reason to assume that corresponding
mechanisms do not apply to fishes and aquatic invertebrates;
• ‘Your eating fuels evolution’
For a long period of time, speciation was thought to only happen between habi-
tats that are geographically isolated. However, the high diversity of fish species
flocks in coral reefs, of cichlid species flocks in Lake Victoria, Lake Tanganyika,
and Lake Malawi, and of sculpin and gammarid species flocks in Lake Baikal ques-
tioned this paradigm and shed light onto food source diversity as a potential evolu-
tionary driver. Food-based bottom-up effects may even lead to sympatric speciation
in fishes and invertebrates (i.e. speciation occurring within the same habitat). This
type of speciation can happen within a few generations. Simply put, this is evolution
in action.
The subject of this book is the nutrition of both wild and farmed finfishes and
aquatic invertebrates, as of course, both farmed aquatic organisms and their wild
counterparts must meet their fundamental needs – replenishing energy and refilling
both inorganic and organic nutrient reserves – through feeding. However, though it
may sound trivial, it must be stated that the scientific approaches and methodologies
of aquaculture and aquatic ecology do differ. Furthermore, theoretical backgrounds
and practical methodologies contrast between the two disciplines in numerous
instances. Throughout the history of science (for more details, refer to Kuhn 2012),
it is well understood that backgrounds and methodologies have a huge impact on the
scientific outcome of an experiment or of a study, even following scientific review.
Therefore, it can be expected that studies either in aquaculture or in the field, even
when concerning identical species, may significantly differ in their results. This is
by no means a drawback. Instead, both aquatic disciplines have a great potential for
scientific cross-fertilization. Aquaculture may learn from aquatic ecology and evo-
lution, whereas ecology may learn from the biochemical and biomolecular path-
ways in individual farmed species.
The background of aquaculture is, according to the Food and Agriculture
Organization (FAO), ‘the farming of aquatic organisms, including fish, mollusks,
crustaceans, and aquatic plants. Farming implies some form of intervention in the
rearing process to enhance production, such as regular stocking, feeding, protection
from predators, etc. Farming also implies individual or corporate ownership of the
stock being cultivated. This definition clearly shows that the focus of aquaculture is
put on individuals of a given species in order to increase production. Relationships
1 Introduction – ‘You Are What You Eat’ 5
to and interactions with the environment are only of interest when identifying and
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reducing potential constraints of production, and keeping it as productive, efficient,

and economical as possible.
In order to avoid endless trial and error, knowledge of the environment, trophic
level, and possible evolutionary relationships of a species is central to understand-
ing the nutritional requirements of that species (this may not be universally appli-
cable however, as not all aquaculture species can be studied in depth in their natural
environment). A striking example of how the ecology of a species can affect its diet,
is the requirement of essential fatty acids in fish species. It is well understood that
the dietary preference for n-3 or n-6 polyunsaturated fatty acids (PUFAs) is deter-
mined by the specific dietary environments. For example, PUFA sources in marine
environments predominantly consist of long-chain n-6 PUFAs. As marine fish
inhabit an environment rich in long chain PUFAs, there is no evolutionary pressure
to retain the ability to endogenously produce long-chain PUFAs by elongation and
desaturation. Conversely, the higher prevalence of shorter PUFAs in freshwaters has
maintained this evolutionary pressure in freshwater fish. Current biomolecular stud-
ies and meta-analyses try to understand the evolution of fatty acid elongation and
desaturation pathways in fishes and invertebrates. This topic will be revisited in
detail in Volume 2.
This PUFA example may be the most notable case of combining dietary require-
ments with the ecological niches of species under consideration. However, there
exists a plethora of examples for so-called trophic bottom-up effects, which demon-
strate that the nutritional basis of lower trophic levels controls the higher ones: for
instance, the diets of herbivores controls the development and well-being of their
predators. A few classical examples may highlight this phenomenon and, further-
more, show that the feedback between ecological niche and species is rather subtle
and even cryptic. Nevertheless, this feedback may serve as a stimulus for aquacul-
turists to study ecological reports more intensively, and vice versa. Crossing the
disciplines of aquaculture and aquatic ecology still has room for improvement, and
could have the potential for new discoveries.
Dietary inorganic bottom-up effects in ecosystems are textbook examples of ter-
restrial and aquatic ecology. Disparities between plant resources and their herbi-
vores in terms of nutrient content not only have major consequences on the success
of the herbivores themselves and their offspring, but also translate to higher trophic
levels and even to ecosystem functioning. For instance, stoichiometric mismatches
between food and consumers affect consumer-driven recycling of limiting nutrients,
which, in turn, functions as a positive or negative feedback on food quality depend-
ing on other sources of nutrient supply. Without doubt, in production-based aqua-
culture, this ecology-based perspective of nutrition is truncated – only one trophic
level is considered.
Food-based bottom-up effects may even lead to sympatric speciation in fishes
and invertebrates within relatively short periods of time. This kind of sympatric
speciation points out the extremely high significance of nutrition to aquatic animals,
though we are only beginning to understand the underlying mechanisms. Doubtless,
genetic variation has often, but not always, been identified as the basis for this kind
of speciation. Alternatively, the action of microRNAs and other epigenetic

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processes are beginning to be taken into consideration. In short-lived animals, such

as protists, nematodes, and fruit flies, it is well documented that epigenetic changes
happen directedly, rather than stochastically, and can be almost as stable as muta-
tions. These animals may serve as guides for future studies with fishes and aquatic
invertebrates, since it can be expected that, by considering diet-driven epigenetic
modifications, at least the broodstocks for aquaculture purposes may be altered and
even improved.
With the still-preliminary total number of marine, brackish, and freshwater fish
species approximated at 29,000 (Lévêque et al. 2008) and the quote ‘A fish is a fish
like a cow is a canary’ from marine biologist Christoffer Schander (Pittman et al.
2013) in mind, I have to confess that ‘Aquatic Animal Nutrition’ can only focus on
a few selected fish and aquatic invertebrate species. Most of those examined are
phylogenetically distant from each other, with inherently wide variations in devel-
opmental strategy. The discrepancy between the number of studied and existing
aquatic invertebrates appears to be even larger than with fishes.
This book does not purely focus on the physiology or ontogenetic development
of fish and aquatic invertebrates, or on increasing productivity in aquaculture.
Numerous textbooks are available that cover these aspects in depth. ‘Aquatic Animal
Nutrition’ comprises two volumes. Volume One presents basic and recent discover-
ies of major feeding mechanisms, as well as long-term effects of diet – from the
individual to the generational level. Volume Two will cover the effects of individual
food ingredients on phenotypic, physiological, and biomolecular levels in the con-
sumers themselves and, where applicable, in their offspring.
Even in two volumes, the following treatise has to be fragmentary and may raise
more issues than fill existing gaps. If, however, the raised issues will initiate new
and innovative studies, this book will have served its purpose.
Appendix
Technical Note
Throughout the books, names and abbreviations of genes are written in lower-case
italics and the abbreviations of the corresponding proteins in capital letters. The
taxonomy of fishes follows ‘fishbase’ and that of invertebrates ‘Encyclopedia of
Life’ and ‘World Register of Marine Species’. If necessary, additional recent revi-
sions were used. Nevertheless, it cannot be guaranteed that no outdated scientific
name has sneaked into this treatise.
References 7
References
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Coad BW, Reist JD (2004) Annotated list of the Arctic marine fishes of Canada. Can Manuscr Rep
Fish Aquat Sci 2674:iv–112
Günther ACLG (1880) An introduction to the study of fishes. Adam & Charles Black, Edinburgh.
https://doi.org/10.5962/bhl.title.54205
Jordan DS (1905) A guide to the study of fishes. H. Holt and Company, New York. https://doi.
org/10.5962/bhl.title.914
Kuhn TS (2012) The structure of scientific revolutions. 50th anniversary, 4th edn. University of
Chicago Press, Chicago
Lévêque C, Oberdorff T, Paugy D, Stiassny MLJ, Tedesco PA (2008) Global diversity of fish (Pisces)
in freshwater. Hydrobiologia 595(1):545–567. https://doi.org/10.1007/s10750-007-9034-0
Pittman K, Yúfera M, Pavlidis M, Geffen AJ, Koven W, Ribeiro L, Zambonino-Infante JL,
Tandler A (2013) Fantastically plastic: fish larvae equipped for a new world. Rev Aquacult
5(SUPPL.1):S224–S267. https://doi.org/10.1111/raq.12034
Chapter 2
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Diets and Digestive Tracts – ‘Your Food

Determines Your Intestine’
Abstract This chapter is an inventory of basic digestive mechanisms and of how

dietary sources determine the forms of the digestive tract of fishes and aquatic inver-
tebrates. Due to high dietary quality and energy provision, carnivores have the
shortest digestive tracts, whereas low dietary quality, low energy, but high fibre
content, determine long intestines of herbivores. However, taken all ecological traits
together, herbivory appears to be a successful, rather than disadvantageous nutri-
tional strategy. The simple relationship of dietary source and intestine length pos-
sesses poor statistical significance and is, therefore, often questioned and apparent
exceptions of this relationship are available. Nevertheless, ontogenetic changes of
intestine length and morphology as well as seasonal changes of the intestine after
dietary alterations support the hypothesis. One major reason for the poor signifi-
cance of this relationship likely are coarse or even incorrect trophic classifications;
consequently, this chapter presents selected examples that species classified as her-
bivores or carnivores are omnivores in reality – the omnivore’s dilemma. Omnivores
have a rather flexible foraging strategy.
2.1 Digestive Tract
To understand the short-term, as well as the long-term and even transgenerational

effects of diets and nutrients, some basic features of the digestive tract have to be
briefly mentioned. This consideration will stay cursory and focus on the dietary
impact on the morphology of the intestine, which can best be observed during onto-
gentic development and metamorphosis, or annual shifts of the trophic niches.
Like other organisms, fishes and aquatic invertebrates require an energy source
to fuel their body systems with fundamental processes, including growth, metabo-
lism and reproduction. Different species have evolved feeding structures and diges-
tive mechanisms that allow them to exploit a vast array of plant and animal food
© Springer Nature Switzerland AG 2018 9

C. E. W. Steinberg, Aquatic Animal Nutrition,
https://doi.org/10.1007/978-3-319-91767-2_2
10 2 Diets and Digestive Tracts – ‘Your Food Determines Your Intestine’
sources. Consequently, the digestive tract of fishes and aquatic invertebrates has
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incorporated numerous adaptations for the efficient breakdown and absorption of

essential nutrients, including appropriate digestive enzymes and absorptive surface
areas. Since the dietary requirements of larvae are different from those of juveniles
or adults, larval nutrition should always be considered along with the organization
and functionality of the digestive system, nutritional needs and the behavior of lar-
vae at different developmental stages. In addition to being the site of nutrient diges-
tion and absorption, the digestive organs provide a barrier to environmental toxins,
confer essential immune function and have important roles in metabolism and salt
and water absorption (Lazo et al. 2011). For more details about the histological and
physiological ontogeny of the digestive tract in fishes, the audience is referred to
this comprehensive, textbook-like presentation.
As a first approximation, the length of the intestine appears to be a function of
the quality and quantity of the available diet (Davis et al. 2013; Karasov and del Rio
2007; Karasov and Douglas 2013; Kramer and Bryant 1995a, b; Wagner et al.
2009). It changes in concert with histological and physiological modifications, sig-
nificantly during the ontogenetic shift from one dietary source to another; for
instance, from insectivory to frugivory, from herbivory to carnivory, or during star-
vation (German et al. (2010); see also Fig. 2.22).
Benavides et al. (1994) presented one of the first experimental demonstrations in
fishes that an increased capability to digest macroalgae is associated with an increase
in relative gut length. This allows larger fish to meet their energetic demands by
consuming algae, owing to their improved capability to digest low-quality food.
Even carnivorous fishes can increase their gut lengths, but herbivores tend to show
a more rapid increase (Kramer and Bryant 1995a). We shall revisit this issue.
Ontogenetic increases in gut length are well known in many, but not all, marine
and freshwater herbivorous fishes (Kramer and Bryant 1995a; Montgomery 1977;
Drewe et al. 2004; Zihler 1981; Ribble and Smith 1983; Stoner and Livingston
1984; Gallagher et al. 2001). More recently, Wagner et al. (2009) reported that diet
quality predicts the intestine length in Lake Tanganyika’s cichlid fishes independent
of their trophic position (Fig. 2.1). The authors tested the effect of trophic positions
on intestine length across 32 species. Trophic positions were inferred from nitrogen
stable isotopes (δ15N), which provide a temporally integrated, quantitative
perspective on the complex diets. Trophic position explains 51% of size-standard-
ized variation in intestine length. Thus, diet is a strong predictor of intestine length
at both intra- and interspecific scales, indicating that fish adjust their phenotype to
balance nutritional needs against energetic costs.
In a meta-study, Karachle and Stergiou (2010) summarized existing literature
data with respect to gut length and Zihler index1 in relationship to the trophic types
(Fig. 2.2). Despite large standard deviations, which will be discussed below, the
trend became clear: herbivores have the longest intestine and carnivores the
shortest.
1
Zihler index (Zihler 1981) is the relation between gut length and body mass: gut
length×(10 × bodymass1/3)−1
2.1 Digestive Tract 11
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Fig. 2.1 Mean size-corrected intestine length and trophic position in the food web (based on mean
δ15N) for 32 species of Lake Tanganyikan cichlids. Regression lines were derived from generalized
least squares regression where (a) λ = 0 (red dashed line; y = 2.4890 ± 0.1640x; r2 = 0.5902),
equivalent to ordinary least squares regression without phylogenetic correction, (b) λ = 0.7180
(ML estimate; black solid line; y = 2.2572 ± 0.1287x; r2 = 0.5093) and (c) λ = 1 (blue dotted line;
y = 2.0371 ± 0.0907x; r2 = 0.4036). Species are color-coded by trophic guild inferred from gut
contents. (From Wagner et al. 2009, courtesy of Wiley)
H H
Functional trophic group
Functional trophic group
OV OV
OA OA
CD CD
CC CC
0 10 20 0 25 50 75
Mean relative gut length Mean Zihler index
Fig. 2.2 Box-plots of mean relative gut length and Zihler’s index values provided by the original
authors for the different functional trophic groups of marine and freshwater fishes, where H herbi-
vores, OA omnivores with preference to animal material, OV omnivores with preference to vegeta-
ble material, CD carnivores with preference to decapods and fish and CC carnivores with preference
to fish and cephalopods. The central box indicates the range of values representing 50% of cases
around the median (vertical lines), the whiskers show the range of the values (horizontal lines), and
cross indicates the mean value (+). (From Karachle and Stergiou 2010, courtesy of the Acta
Ichthyologica et Piscatoria) (Although the authors use trophic classifications, the risk of pure nom-
inal classifications or even misclassifications cannot be excluded; see also ‘Trophic Positions: An
Omnivores’ Dilemma?’ below. Nominal, rather than empirically observed, classifications surely
increase the standard deviation.)
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Fig. 2.3 Relationship between phylogenetically independent contrasts of intestinal length residu-
als and contrasts of arcsine transformed proportion of animal material in diet. Numbers refer to
different grunter species. (From Davis et al. 2013, courtesy of Biomed Central Ltd.)
So far, the relationship between trophic position and length of the intestine and
its evolutionary derivation have received little attention from a phylogenetic per-
spective. Davis et al. (2013) documented the phylogenetic development of intestinal
length variability, and resultant correlation with dietary habits, within a molecular
phylogeny of 28 species of terapontid fishes. They found that the shorter the
intestine, the higher the share of animal preys in the diet (Fig. 2.3). The Terapontidae
(grunters), an ancestrally euryhaline-marine group, is the most trophically diverse
of Australia’s freshwater fish families, with widespread shifts away from
animal-prey-dominated diets occurring since their invasion of freshwaters. The
ontogenetic development of intestinal complexity appears to represent an important
functional innovation underlying the extensive trophic differentiation, specifically
facilitating the pronounced shifts away from the carnivorous (including inverte-
brates and vertebrates) diets evident across the family. The capacity to modify intes-
tinal morphology and physiology appears to be an important facilitator of trophic
diversification during the phyletic radiations – not only within the grunters.
Several striking examples prove that the intestine length varies in a single spe-
cies, depending on the available diet source. Investigating how the diet and intesti-
nal length of a persistent and generalist fish species (Bryconamericus iheringii,
Characidae) responds to riparian modifications in 31 subtropical streams in south-
ern Brazil, Dala-Corte et al. (2017) showed that the generalist and locally persistent
fish species responded to environmental alterations caused by riparian degradation
2.1 Digestive Tract 13
by consuming a greater proportion of autochthonous material (algae and aquatic

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macrophytes) instead of allochthonous material (terrestrial plant fragments and ter-

restrial invertebrates). These findings indicate that plasticity in intestinal length is an
important characteristic to determine whether fish populations can persist in a vari-
ety of habitat conditions and cope with the digestion of a greater proportion of low-
quality and low-protein food items in human-altered environments.
The intestine length also varies on an annual basis. In the Trinidadian guppy,
Zandona et al. (2015) showed that, even in omnivorous fish, gut length adapted to
different diets, being more evident when the magnitude of difference between ani-
mal and plant material in the diet was very large. The authors sampled guppies from
sites with low (LP) and high predation (HP) pressure in the Aripo and Guanapo
Rivers in Trinidad. They collected fish during the dry and wet seasons and assessed
their diet and gut length. During the dry season, guppies from HP sites fed mostly
on invertebrates, while guppies in the LP sites fed mainly on detritus. During the
wet season, the diet of LP and HP populations became very similar.
This study indicates that guppies have a broad range of variation in the propor-
tion of invertebrates and detritus in their diet, which changed with season and was
associated to local adaptation. These variations in diet were correlated with the gut
length (Fig. 2.4). Guppies that showed higher levels of carnivory also had the
Fig. 2.4 Mean proportion of invertebrates in diets vs. mean relative gut length. Only guppies
between 14 and 20 mm were included. Each data point represents one site (Aripo HP and LP for
both dry and wet season, and Guanapo HP and LP from the dry season). Relative gut length was
calculated as the gut length divided by fish length. An average value was assigned for the propor-
tion of invertebrates for each site, which was the estimated marginal mean obtained from the diet
analysis. (From Zandona et al. 2015, courtesy of the Public Library of Science)
shortest guts, and vice versa, as those with higher levels of herbivory had longer
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guts. The flexibility of the digestive system is an important adaptation that enables
guppies (and other fish species) to respond favorably to changes in food sources and
maximize nutrient absorption and energy extraction from different food types.
Different gut types can also be observed during the ontogenetic development, as
displayed, for instance, in Brycon guatemalensis (Drewe et al. 2004), a Neotropical
characid fish. It consumes an entirely terrestrial diet, shifting from eating insects as
juveniles to fruits and leaves as adults. Juveniles and larger-sized fishes were stud-
ied to test the hypotheses that, with ontogeny, (1) relative gut length increases, (2)
pyloric ceca arrangement and number remain unchanged and (3) pepsin, trypsin and
lipase activities decrease, while α-amylase activity increases. These hypotheses
were supported in that larger fish had longer guts, unchanged pyloric ceca arrange-
ment, and lower pepsin and trypsin activities, but higher α-amylase activities than
the juveniles. This study supports the view that B. guatemalensis is specialized mor-
phologically and biochemically to function first as a carnivore and then as an herbi-
vore during its life history.
2.2 Digestion
One major function of the intestine is digestion. This is the process of hydrolysis
and solubilization of ingested nutrient polymers into molecules and elements suit-
able for transport across the intestinal wall. The digestive enzymes secreted from
the stomach and exocrine pancreas are of major importance for enzymatic hydroly-
sis of complex food polymers, such as proteins, fats and carbohydrates, into smaller
fragments. The resulting smaller fragments are further digested at the epithelium of
the intestinal tract by the enzymes located in the brush border membrane of the
enterocytes, releasing molecules small enough for absorption, i.e. small peptides
and amino acids, monosaccharides, and fatty acids. This process is summarized in
Fig. 2.5. However, the contribution of exogenous digestive enzymes present in the
natural diet to total digestive capacity has most likely been largely underestimated.
A recent review focuses on exogenous contributions to digestion in fishes (see
Kuz’mina (2008) and Functional Aquafeed, Volume 2) and the following will there-
fore focus on endogenous gastrointestinal digestion processes. Considering the
importance of providing cultured fish with highly digestible formulated feeds for
rapid, cost-efficient fish growth and low waste released to the environment, the vast
majority of the investigations on digestive processes and factors that affect nutrient
digestibility have been carried out on production fish.
Fish have a digestive enzyme apparatus qualitatively similar to that of other ani-
mals with very similar substrate specificities across taxonomic groups. Although
molecular characterizations are now being published with increasing frequency,
knowledge is still limited regarding more specific characteristics of various
digestive enzymes for most fish species. Species-specific isoforms of the various
enzymes exist with differences in, for example, molecular-weights, specific activi-
2.2 Digestion 15
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Fig. 2.5 Schematic drawing of the digestive processes along the digestive tract of fish. The loca-
tion of various enzymes and other digestive components and the respective processes in the lumen,
as opposed to the intestinal mucosa, are indicated. FFA free fatty acids, FSVit fat soluble vitamins
Design: F. Venold. (From Bakke et al. 2010 with permission from Elsevier)
ties, pH-optima and efficiencies towards different bonds. Fish enzymes typically
show higher specific activity and substrate affinities than those in homeothermic
animals, presumably representing an evolutionary adaptation to function at lower
temperatures. For example, trypsin from Atlantic cod has a 17-times higher catalytic
efficiency than bovine trypsin when measured at the same temperature range.
2.2.1 Protein Digestion
In fish species with stomachs, the low pH from HCl secretion denatures most of the
proteins as they are solubilized, opening the structure for easier access by the pro-
teolytic enzyme pepsin. Pepsinogen and pepsin from several fish species have been
characterized. The enzyme is present in fishes in more than one form, and the dif-
ferent forms show different activation rates, pH optima (varying between 1 and 5),
specific activities and substrate specificities. Pepsins are endopeptidases, i.e. they
hydrolyze peptide bonds, with a high affinity for hydrophobic bonds involving
amino acids (AAs), such as tyrosine (Tyr) and phenylalanine (Phe). The partial
hydrolysis of the proteins increases the solubility and dissolution of other food com-
ponents, and prepares the diet—after this stage called chyme—for entry into the
intestine through the pyloric sphincter.
Proteins and peptides entering the intestine, with or without prior processing in a
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stomach, are diluted and dissolved in alkaline secretions from the liver, pancreas
and/or gut wall. The actions of the pancreatic endopeptidases trypsin, chymotrypsin
and elastases I and II, as well as the exopeptidases carboxypeptidase A and B, result
in a mixture of free AAs and smaller peptides (Fig. 2.5). The final steps of peptide
hydrolysis take place at the brush border of the enterocytes by aminopeptidases, or
by intracellular peptidases following peptide transport across the membrane.
However, some proteins and peptides entering the intestine either from the diet,
gastrointestinal or pancreatic secretions, may resist proteolysis and reach the distal
intestine more or less intact.
2.2.2 Lipid Digestion
Efficient lipid digestion requires emulsifiers in the mixture of food—mainly pro-

teins and phospholipids—as well as from endogenous bile acid and phospholipid
secretion in the proximal part of the digestive tract. The emulsifiers orient them-
selves on the surface of lipid droplets that form as dietary lipid is released during the
physical, chemical and enzymatic degradation of the food. If the emulsifying capac-
ity is deficient, the digestion of released lipids may be hindered. The main source of
lipolytic enzymes in fish is the acinar cells of theexocrine pancreas. Lipase activity
differs between fish species, as illustrated by the difference between related species,
such as Atlantic salmon and rainbow trout. Active fishes such as mackerel (family
Scombridae) and scup (family Sparidae) are among the species that have especially
high activities. Knowledge of characteristics and specificities of fish lipases is far
from complete. Freshwater fishes may have mainly co-lipase-dependent pancreatic
lipase (PL), whereas marine fishes have bile-acid-dependent carboxyl ester lipase
(CEL). PL has higher specificity and digestive efficiency for triglycerides than the
CEL. The latter hydrolyses a broader range of lipids, including wax esters (Bakke
et al. (2010) with reference).
2.2.3 Carbohydrate Digestion
Carbohydrates in natural fish diets and formulated feeds range from the highly sol-
uble and digestible mono-, di- and oligosaccharides, glycogen and starch, to only
marginally soluble and digestible chitin, hemicelluloses and celluloses. Fish species
vary greatly in their capacity to digest and absorb even soluble carbohydrates. Some
may have developed intestinal structures, functions and microbiota that enable
hydrolysis of a greater variety of carbohydrates, although this appears to be variable
even among herbivorous species. Fish have two categories of endogenous enzymes
2.3 Ontogenesis and the Intestine 17
involved in carbohydrate digestion: pancreatic α-amylase and disaccharidases in the

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brush border membrane of the intestinal epithelial cells (Fig. 2.5).
2.3 Ontogenesis and the Intestine
2.3.1 F
ishes
According to the classical and illustrative paper by Dabrowski (1984), the ontoge-
netic changes in digestive tract development of freshwater fishes during the larval–
juvenile transition can be categorized into three types:
1. Stomachless fish with an increase in complexity of the coiling pattern (mainly
cyprinids) (Fig. 2.6). These fishes remain stomachless throughout life. However,
in this group the coiling pattern of the intestine undergoes ontogenetic changes;
2. Stomachless larvae which develop a stomach structure after ingestion of food
(coregonids, silurids, serrasalmids) (Fig. 2.7); and
3. Alevin and juvenile stages of fish capable of ingesting the first food when the
stomach is present as a distinguished feature (salmonids, cichlids) (Fig. 2.8). In
other words, salmonids appear to have a functional stomach, before changing
from endogenous to external food. In the ontogeny of the cichlid digestive tract,
the small stomach is visible before yolk-sac absorption and, as the fishes take
their first external food, the stomach appears as a sizeable blind pouch. In spite
of the extremely different feeding habits of this species group (algae, plants,
fruits, detritus, insects, or fish), the overall appearance of the digestive tract
remains the same.
Morphological features of the digestive system are of great consequence in
respect to the diet type that larval/juvenile fish are able to utilize, especially at the
high growth rates during early ontogenetic development [50% per day in larval
common carp (Cyprinus carpio), 30–50% per day in the African catfish (Clarias
gariepinus) larvae]. Cichlids are exceptional, as their digestive gastrointestinal tract
appears to be completely formed with a functional stomach and an elongated intes-
tine prior to the use of yolk sac reserves. Unlike most other teleosts, cichlid juveniles
pass through an extended period of “mixed” feeding of endogenous (yolk sac) and
exogenous feeding. This modulation shifts the focus to maternal–offspring nutrient
transfer in juveniles, rather than a sole dependence on external food intake and its
quality (nutrient presence and availability) for larval fish. Juvenile, first feeding Nile
tilapia, for instance, were able to grow on phytoplankton (especially, coccal green
algae) provided during the first several weeks of life (Dabrowski and Portella
(2005)).
In addition to the well-documented freshwater species, Fig. 2.9 exemplifies the
ontogenetic development of the digestive system in the orange clownfish, a species
Swim bladder inflates

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Trypsin activity rises

exogenous feeding
Chymotrypsin activity appears

weight (mg)
Amino peptidase activity increases
1000
100
10
1
15 30
age (days)
Fig. 2.6 Ontogenetic development of the cyprinid fish digestive tract, exemplified by common
carp. (From Dabrowski 1984, courtesy of Editions scientifiques medicales Elsevier, Paris)
with an advanced alimentary canal at hatching. These larvae can immediately start
exogenous feeding.
A second marine example, Elbal et al. (2004) reported light and electron micro-
scopic studies of the digestive tract of the gilthead sea bream (Sparus aurata,
Fig. 2.10) from hatching to 69 days. Five significant phases were established. Phases
I and II comprise the lecitotrophic period. During phase I, the yolk sac was large
and the uniform digestive tract showed a layer of squamous epithelial cells with
Exogenous feeding
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Trypsin + chymotrypsin activity present

Guanine in skin (metamorphosis)
weight (mg)
Glyco-glycogeno-lysis as energy

Morphologically functional
stomach
Stomach acid pH
pepin digestion
150
100
50
10
30 60
age (days)
Fig. 2.7 Ontogenetic development of the corregonid fish digestive tract, exemplified by Coregonus
pollan. (From Dabrowski 1984, courtesy of Editions scientifiques medicales Elsevier, Paris)
numerous free ribosomes. Phase II was characterized by the opening of the anus and
the differentiation of three digestive regions: the esophagus, with a stratified epithe-
lium; the presumptive stomach, whose cuboid epithelial cells had some apical pro-
cesses and clear vesicles; and the intestine, with large intercellular spaces among
prismatic epithelial cells that had a periodic acid Schiff reagent-positive striated
border. Phase III, or lecitoexotrophic period, began with the opening of the mouth,
where absorption of the yolk sac started and the intestine became differentiated into
two regions separated by a valve. Intestinal epithelial cells showed basal lamellar
structures and lipoprotein particles. Some columnar cells appeared inside the epi-
thelium of the esophagus. Phases IV and V comprise the exotrophic period, where
phase IV begins with the disappearance of the yolk sac; mucous cells containing
Functional pancreas and gastric glands

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Exogenous feeding
Trypsin activity increases

weight (mg)
200
100
30 60
age (days)
Fig. 2.8 Ontogenetic development of the salmonid fish digestive tract, exemplified by rainbow
trout. (From Dabrowski 1984, courtesy of Editions scientifiques medicales Elsevier, Paris)
sulphomucin-type acid mucosubstances appeared in the esophagus and goblet cells,

with acid and neutral mucosubstances appeared in the intestine. The epithelial cells
of the first and posterior intestinal segments showed large lipid droplets and heavy
pinocytosis, with large supranuclear vesicles and numerous lysosomes, respectively.
Phase V was marked by the appearance of neutral mucosubstances in the esopha-
geal mucous cells and in the stomach epithelial cells, and the differentiation of
pyloric ceca and gastric glands. The ultrastructural features of glandular cells indi-
cated that they secrete both pepsinogen and hydrochloride acid. The epithelial cells
of the first intestinal segment showed large lipid droplets, often close to mitochon-
dria, at the beginning of this phase. These lipid droplets decreased in size, while the
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Juvenile
Growth and Differentitation
Metamorphosis
e
Digestion becomes
siz extracellular
n
s i ar
mn cell der
s
se mn Larva
gastric gland
colu ach bor
a
s
re olu
inc s c
sh
en cell
ru
ar
lum gut tb
gu
m
d
sto
nd
hin
Hi
Hatching
Mouth opened Onset of exogenous feeding
Jaws ossified Digestion is pinocytotic Embryo
Alimentary canal
differentiated
Liver and spleen
differentiated
Six days 1 3 5 5 5 7
after
fertilisation Age (Days after hatch)
Fig. 2.9 Left: Steps in the ontogeny of the digestive system of the orange clownfish (Amphiprion
percula). The various structures of the alimentary canal grow and differentiate at different rates,
but are completed and functional at the same time, enabling the larvae to undergo rapid metamor-
phosis from one stage to another. The steps in A. percula development have been divided into
embryonic, larval, and juvenile stages. There is no eleutheroembryo stage as the larvae begin
exogenous feeding immediately after hatching and before the yolk sac is fully absorbed. (From
Gordon and Hecht 2002, with permission from Wiley); right: Breeding A. percula (courtesy of
Haplochromis, Wikimedia)
Fig. 2.10 Images of Sparus aurata and Gadus morhua. (From Bloch 1785–1790, courtesy of the
Biodiversity Heritage Library)
rough and smooth endoplasmic reticulum and Golgi complex, related to lipoprotein
synthesis, progressively developed during this phase. Pinocytotic and large supra-
nuclear vesicles disappeared from epithelial cells of the posterior intestinal
segment.
Overall, in the early development stages of S. aurata, lipid absorption occurs in
the epithelial cells of the first intestinal segment, while the absorptive cells of the
posterior intestinal segment are able to take up proteins by pinocytotic mechanisms.
The appearance of the first gastric glands improves extracellular protein digestion,
and the supranuclear inclusions in the absorptive cells of the posterior intestinal
segment disappear. This event probably encourages lipoprotein formation and the
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exportation of lipids in the form of chylomicrons. These findings are of considerable

importance for the evaluation of the digestive tract functionality, and will be useful
for establishment of optimal rearing techniques and artificial food for S. aurata
larvae for the commercial production of this species.
Pedersen and Falk-Petersen (1992) compared the development of Atlantic cod
(Gadus morhua, Fig. 2.10) during transition from larva to juvenile with that of four
other teleosts, namely red seabream (Pagrus major), yellowtail (Seriola quinquera-
diata), spot croaker (Leiostomus xanthurus) and olive flounder (Paralichthys oliva-
ceus) (Fig. 2.11). The differentiation of the alimentary tract during the early stages
of juvenile development, into a stomach originating from the posterior part of the
esophagus, and pyloric ceca2 developing from the anterior part of the intestine, is in
accordance with the general scheme seen in other teleosts. In cod, the developmen-
tal changes in the alimentary tract occur at a later stage than the disappearance of
the median finfold, the development of the median fins and the appearance of the
vertebrae. Compared with some other fish species, alimentary tract development
appears to be delayed in cod. In red sea-bream, spot and olive flounder, the stomach
and pyloric ceca develop at the time of the disappearance of the median finfold
(Fig. 2.11). In yellowtail and cod, the development of stomach and pyloric ceca
takes place later than the development of characters associated with locomotion
(Fig. 2.11). In cod, the food-storing capacity of the stomach seems to develop in
individuals larger than 20 mm and this capacity is well-developed in individuals
larger than 40 mm. In fish of 40 mm and over, the vast majority of identifiable cope-
pods in the alimentary system were found in the stomach.
The strategy of expanding intestinal absorptive surface is taking place in both
cold water and tropical fishes. In the cold-water salmonid rainbow trout, which pos-
sesses on average 70 pyloric ceca, the total length of the pyloric ceca is 6-fold larger
than the total intestine. Several tropical characid fishes also possess numerous
pyloric ceca. A generalization predicts that diets composed of voluminous food,
algae and detritus should result in elongation of the intestine. Thus, the relative
length of the intestine seems to be an indicator of diversified use by the digestive
tract of food of high or low nutrient concentrations (Fig. 2.12, Dabrowski and
Portella (2005)). For instance, the Lake Tanganyika cichlid Petrochromis polyodon
possesses a particularly long intestine (6–10-fold body length), because it ingests
unicellular microalgae by scraping the rock biocover (Takamura 1984). In contrast
to this fish species, the carnivorous channel catfish (Ictalurus punctatus) is charac-
terized by the shortest relative intestine length (Fig. 2.12). Its diet consists primarily
of small fish, crustaceans (e.g. crayfish), clams and snails; it feeds, also, on aquatic
insects, even small mammals, but seldom plant material (Tyus and Nikirk 1990).
The concept of nutrient requirement and morphology of the intestine has been
questioned several times. For instance, there are strict stomachless carnivorous fish
2
Pyloric ceca are blind appendages attached to the proximal intestine of many fish. Buddington
and Diamond (1987) provided evidence that ceca are an adaptation for increasing the intestinal
surface area without increasing the length or thickness of the intestine itself.
Species Standard length (mm)

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character 0 10 20 30 40 50
Gadus morhua1
eleutheroembryo I
median finfold
median fins
stomach
pyloric caeca
Pagrus major 2
eleutheroembryo I
median finfold
median fins
stomach
pyloric caeca
gastric glands
Seriola quinqueradiata 3
eleutheroembryo I
median finfold
median fins
stomach
pyloric caeca
gastric glands
Leiostomus xanthurus 4
eleutheroembryo I
median finfold
median fins
stomach
pyloric caeca
Paralichthys alivaceus 5
eleutheroembryo I
median finfold
median fins
stomach
pyloric caeca
Fig. 2.11 The development scheme of Atlantic cod (Gadus morhua) during transition from larva
to juvenile, compared with those of some other teleost species. (From Pedersen and Falk-Petersen
1992, with permission from Wiley)
(the cyprinid Leuciscus aspius) and herbivores with short digestive tract, 1.5 × body
length (Horn 1989). Another example underlines this concern, but does not suspend
the general trend displayed in Fig. 2.1 and in the meta-study referred to in Fig. 2.2.
Rather, it contributes to the understanding of the huge standard deviations in the
latter figure. In particular, German and Horn (2006) measured relative gut length,
body mass (Zihler’s index) and relative gut mass in four species of prickleback
fishes, a group of fish which will serve as witness for the considerations about
herbivory.
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Fig. 2.12 Changes in the relative length of intestine (expressed in body lengths) in several fish
species. (From Dabrowski and Portella 2005, with permission from Elsevier)
The authors were interested in the effects of ontogeny, diet and phylogeny on
these gut dimensions (Fig. 2.13). Of the four species, Cebidichthys violaceus and
Xiphister mucosus shift to herbivory with growth, whereas X. atropurpureus and
Anoplarchus purpurescens remain carnivores. A. purpurescens belongs to a carniv-
orous clade, and the three other species belong to an adjacent, herbivorous clade.
The comparison of the gut dimensions in three feeding categories of the four species
revealed:
1. Small, wild-caught juveniles representing the carnivorous condition before two
species shift to herbivory;
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Fig. 2.13 Upper graph: Non-metric multidimensional scaling plot of all three gut dimension
parameters combined for each species as a function of ontogeny (w30–40 and W60–75 categories – the
subscripts refer to body lengths in mm) in Cebidichthys violaceus (Cv), Xiphister mucosus (Xm),
X. atropurpureus (Xa) and Anoplarchus purpurescens (Ap). Arrows indicate magnitude of ontoge-
netic shifts in gut dimensions. The stress value indicates that the plot fits well (i.e., values <0.1)
into two-dimensional space. Lower graph: Non-metric multidimensional scaling plot of all three
gut dimension parameters combined for each species as a function of diet (W60–75 and L60–75 catego-
ries) in C. violaceus (Cv), X. mucosus (Xm), X. atropurpureus (Xa) and A. purpurescens (Ap).
Arrows indicate magnitude of diet-related changes in gut dimensions. The stress value indicates
that the plot fits well (i.e., values <0.1) into two-dimensional space. (From German and Horn 2006,
with permission from Springer)
2. Larger, wild-caught juveniles representing the natural diet condition of the two
carnivores and the two species that have shifted to herbivory; and
3. Larger, laboratory raised juveniles produced by feeding an artificially high-
protein diet to small juveniles until they have reached the size of the larger, wild-
caught juveniles.
Comparisons of the gut dimensions in categories (1) versus (2) tested for an
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ontogenetic effect, in (2) versus (3) for a dietary effect, and within each category for
a phylogenetic effect. C. violaceus and X. mucosus increased gut dimensions with
an increase in body size, and did not change the ontogenetic trajectory in gut dimen-
sions on the artificially high-protein diet, indicating that they are genetically pro-
grammed to develop relatively large guts associated with herbivory. X. atropurpureus
increased its gut dimensions with an increase in size similar to its sister taxon, X.
mucosus, indicating a phylogenetic influence, but decreased gut dimensions on the
artificially high-protein diet, showing phenotypic plasticity. Nevertheless, X. atro-
purpureus displayed a larger gut than A. purpurescens, further evidence that it
evolved in an herbivorous clade. A. purpurescens possessed a relatively small gut
that was less affected by ontogeny or diet. Overall, ontogeny and phylogeny, more
than diet, appear to influence gut dimensions in the four species, thus favoring
genetic adaptation over phenotypic plasticity as the major force acting on digestive
system features in the two prickleback clades. This contrasts the metastudy in
Fig. 2.2.
Further details of ontogeny and physiology of the digestive system of marine and
freshwater fish larvae and adults can be found in recent monographs and reviews
(Bucking 2015; Zambonino Infante et al. 2008; Portella and Dabrowski 2008;
Yúfera and Darias 2007; Zhang et al. 2017; Grosell et al. 2010; Cortés et al. 2008).
2.3.2 Invertebrates
Dietary impact on the development of the intestines in aquatic invertebrates is much

less well documented than in fish species. We shall focus on selected examples from
echinoderms and crustaceans.
2.3.2.1 E
chinoderms
Sea urchins exhibit dramatic changes in the ontogenetic niche between newly set-
tled recruits, that may occupy cryptic habitats and feed on microalgae or detritus,
and mature adults, that may catch drift kelp or actively graze on macroalgae and
invertebrates. Wing and Wing (2015) examined patterns in the ontogenetic niche
development of Evechinus chloroticus within the New Zealand fjords, a system with
a series of strong environmental and benthic productivity gradients. E. chloroticus,
also known as “kina” (from the Māori name), is a sea urchin endemic to New
Zealand (Fig. 2.14). Using data from 15 long-term monitoring sites, the authors
examined relationships between the diet of juvenile and adult sea urchins and the
morphology of their preferred food, the kelp Ecklonia radiata. Stable isotope analy-
sis (δ13C and δ15N) of E. chloroticus stomach contents, muscle tissue and samples
along the blades of E. radiata revealed evidence of large variation in nutritional
conditions. The contribution of E. radiata to the diet of E. chloroticus declined from
Another random document with
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The Project Gutenberg eBook of Ironheart
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States and most other parts of the world at no cost and with
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Title: Ironheart
Author: William MacLeod Raine
Release date: November 6, 2023 [eBook #72057]
Language: English
Original publication: New York: Grosset & Dunlap, 1923
Credits: Tim Lindell, David E. Brown, and the Online Distributed

Proofreading Team at https://www.pgdp.net (This file
was produced from images generously made available
by The Internet Archive/American Libraries.)
*** START OF THE PROJECT GUTENBERG EBOOK IRONHEART

***
IRONHEART
IRONHEART
BY
WILLIAM MacLEOD RAINE
AUTHOR OF
MAN-SIZE, TANGLED TRAILS,
THE FIGHTING EDGE, Etc.
GROSSET & DUNLAP

PUBLISHERS NEW YORK
Made in the United States of America

COPYRIGHT, 1922, BY THE CURTIS PUBLISHING COMPANY
COPYRIGHT, 1923, BY WILLIAM MACLEOD RAINE
ALL RIGHTS RESERVED
PRINTED IN THE U.S.A.

TO
ARTHUR CHAPMAN
“But somehow when he’s gone, you think a heap
About his virtues—how he’s square and true;
If more come stringin’ in they’d make it cheap—
This friendship thing—and spoil it all for you.”
CONTENTS
I. Turfing it 3
II. “De King o’ Prooshia on de Job” 8
III. One of the Lost Legion 15
IV. Betty rides 19
V. Tug is “collected” 31
VI. “Nothing but a Gay-Cat anyhow” 35
VII. Tug says, “No, Thank You” 43
VIII. A Rift in the Lute 48
IX. Under Fire 55
X. “One Square Guy” 65
XI. Mr. Ne’er-do-Well 72
XII. “Is this Bird a Prisoner, or ain’t he?” 80
XIII. A Job 85
XIV. One Bad Hombre meets Another 89
XV. The Homesteader serves Notice 98
XVI. The Stampede 105
XVII. His Picture in the Paper 111
XVIII. A Hot Trail 116
XIX. Captain Thurston K. Hollister 122
XX. A Clash 130
XXI. Irrefutable Logic 142
XXII. A Stern Chase 147
XXIII. Out of the Blizzard 157
XXIV. “Come on, you Damn Bushwhacker” 166
XXV. A Difference of Opinion 174
XXVI. Black is Surprised 182
XXVII. The Man with the Bleached Blue Eyes 189
XXVIII. Betty has her Own Way 196
XXIX. A Child of Impulse 202
XXX. Fathoms Deep 209
XXXI. Betty makes a Discovery 214
XXXII. Without Rhyme or Reason 224
XXXIII. The Bluebird alights and then takes Wing 230
XXXIV. Born that Way 237
XXXV. Birds of a Feather 246
XXXVI. A Stormy Sea 252
XXXVII. Hold the Fort 262
XXXVIII. Beyond a Shadow of a Doubt 269
XXXIX. The Turn of a Crooked Trail 277
XL. Betty discovers why she is Young 284
IRONHEART
IRONHEART
CHAPTER I
TURFING IT
A thin wisp of smoke drifted up from the camp at the edge of the
wash. It rose languidly, as though affected by the fact that the day
was going to be a scorcher. Already, though the morning was young,
a fiery sun beat down on the sand so that heat waves shimmered in
the air. Occasionally a spark from the crackling cottonwood limbs
was caught by a dust whirl and carried toward the field of ripe wheat
bordering the creek.
Of the campers there were three, all of the genus tramp, but each a
variant. They represented different types, these desert trekkers.
The gross man lying lazily under the shade of a clump of willows
might have stepped straight out of a vaudeville sketch. He was dirty
and unkempt, his face bloated and dissipated. From his lax mouth
projected an English brier pipe, uncleansably soiled. His clothes
hung on him like sacks, wrinkled and dusty, but not ragged. He was
too good a hobo to wear anything torn or patched. It was his boast
that he could get another suit for the asking any time he needed one.
“I’m a blowed-in-the-glass stiff,” he bragged now. “Drilled from
Denver to ’Frisco fifteen times, an’ never was a stake man or a
shovel bum. Not for a day, ’boes. Ask any o’ the push about old York.
They’ll give it to youse straight that he knows the best flops from
Cincie to Phillie, an’ that no horstile crew can ditch him when he’s
goin’ good.”
York was a hobo pure and simple. It was his business in life. For
“stew-bums” and “gay-cats,” to use his own phraseology, he had a
supreme contempt. His companions were amateurs, from his point of
view non-professionals. Neither of them had any pride in turfing it,
which is the blanket stiff’s expression for taking to the road. They did
not understand York’s vocabulary nor the ethics that were current in
his craft.
Yet the thin, weasel-faced man with the cigarette drooping from his
mouth was no amateur in his own line. He had a prison face, the
peculiar distortion of one side of the mouth often seen in confirmed
criminals. His light-blue eyes were cold and dead. A film veiled them
and snuffed out all expression.
“Cig” he called himself, and the name sufficed. On the road
surnames were neither asked for nor volunteered. York had sized
him up three days before when they had met at Colorado Springs,
and he had passed on his verdict to the third member of the party.
“A river rat on a vac—hittin’ the grit for a getaway,” he had
whispered.
His guess had been a good one. Cig had been brought up on the
East River. He had served time in the penitentiaries of three States
and expected to test the hospitality of others. Just now he was
moving westward because the East was too hot for him. He and a
pal had done a job at Jersey City during which they had been forced
to croak a guy. Hence his unwilling expedition to the Rockies. Never
before had he been farther from the Atlantic than Buffalo, and the
vast uninhabited stretches of the West bored and appalled him. He
was homesick for the fetid dumps of New York.
The corner of his mouth lifted in a sneer. “Wot’ell would any one
want to cross this Gawd-forsaken country fifteen times for unless he
was bughouse?”
“If you read the papes you’d know that travel is a lib’ral ejucation.
Difference between a man an’ a tree is that one’s got legs to move
around with. You ginks on the East Side act like you’re anchored to
the Batt’ry an’ the Bowery. Me, I was born there, but I been batterin’
on the road ever since I was knee-high to a duck. A fellow’s got to
throw his feet if he wants to learn,” York announced dogmatically.
There was obvious insult in Cig’s half-closed eyes. “’S at what
learned you all youse know?” he asked.
“Don’t get heavy, young feller,” advised the blanket stiff. “I’ve knew
guys to stay healthy by layin’ off me.”
The young man cooking breakfast barked a summons. “Come an’
get it.”
The tramps moved forward to eat, forgetting for the moment their
incipient quarrel. Into tin cups and plates the cook poured coffee and
stew. In his light clean build, slender but well-packed, was the
promise of the athlete. His movements disappointed this expectation.
He slouched, dragging the worn shoes through the cracks of which
the flesh was visible. Of the three, he was the only one that was
ragged. The coat he wore, which did not match the trousers, was at
the last extremity.
One might have guessed his age at twenty-three or four. If it had not
been for the sullen expression in the eyes and the smoldering
discontent of the face, he might have been good-looking. The
reddish hair was short, crisp, and curly, the eyes blue as the
Colorado summer sky above, the small head well-shaped and
beautifully poised on the sloping column of the neck.
Yet the impression he made on observers was not a pleasant one.
His good points were marred by the spirit that found outlet in a sullen
manner that habitually grudged the world a smile. He had the skin
pigment of the blond, and in the untempered sun of the Rockies
should have been tanned to a rich red-brown. Instead, the skin was
clammily unhealthy. The eyes were dulled and expressionless.
York ate wolfishly, occasionally using the sleeve of his coat for a
napkin. He talked, blatantly and continuously. Cig spoke only at rare
intervals, the cook not at all. Within the silent man there simmered a
nausea of disgust that included himself and all the universe in which
he moved.
In the underworld caste rules more rigidly than in upper strata of
society. The sense of superiority is everywhere an admission of
weakness. It is the defense of one who lives in a glass house. Since
all of these men were failures, each despised the others and
cherished his feeling that they were inferior.
“You ’boes turf it with York an’ you’ll always have plenty o’ punk and
plaster,” the old tramp swaggered. “Comes to batterin’ I’m there with
bells on.”
Translated into English, he meant that if they traveled with him they
would have bread and butter enough because he was a first-class
beggar.
“To hear youse chew the rag you’re a wiz, ain’t you?” Cig jeered. “I
ain’t noticed you diggin’ up any Ritz-Carlton lunches a guy can write
home about. How about it, Tug?”
The cook grunted.
“Me, I can tell a mark far as I can see him—know whether he’s good
for a flop or a feed,” York continued. “Onct I was ridin’ the rods into
Omaha—been punchin’ the wind till I was froze stiff, me ’n’ a pal
called Seattle. Shacks an’ the con tried to ditch us. Nothin’ doing. We
was right there again when the wheels began to move. In the yards
at Omaha we bumps into a gay-cat—like Tug here. He spills the
dope that the bulls are layin’ for us. Some mission stiff had beefed
on me. No guy with or without brass buttons can throw a scare into
old York. No can do. So I says to Seattle, says I—”
York’s story died in his throat. He stood staring, mouth open and chin
fallen.
Two men were standing on the edge of the bluff above the bed of the
creek. He did not need a second look to tell him that they had come
to make trouble.
CHAPTER II
“DE KING O’ PROOSHIA ON DE JOB”
To Reed came his foreman Lon Forbes with a story of three tramps
camping down by Willow Creek close to the lower meadow
wheatfield.
The ranchman made no comment, unless it was one to say, “Get out
the car.” He was a tight-lipped man of few words, sometimes grim.
His manner gave an effect of quiet strength.
Presently the two were following the winding road through the
pasture. A field of golden wheat lay below them undulating with the
roll of the land. Through it swept the faintest ripple of quivering grain.
The crop was a heavy one, ripe for the reaper. Dry as tinder, a spark
might set a blaze running across the meadow like wildfire.
Forbes pointed the finger of a gnarled hand toward a veil of smoke
drifting lazily from the wash. “Down there, looks like.”
His employer nodded. They descended from the car and walked
along the edge of the bank above the creek bed. Three men sat near
a camp-fire. One glance was enough to show that they were hoboes.
Coffee in an old tomato can was bubbling over some live coals set
between two flat stones.
The big man with the bloated face was talking. The others were
sulkily silent, not so much listening as offering an annoyed refusal to
be impressed. The boaster looked up, and the vaporings died within
him.
“What you doing here?” demanded Reed. His voice was curt and
hostile.
York, true to type, became at once obsequious. “No offense, boss. If
these here are private grounds—”
“They are,” the owner cut in sharply.
“Well, we’ll hit the grit right away. No harm done, mister.” The voice
of the blanket stiff had become a whine, sullen and yet fawning.
His manner irritated both of his companions. Cig spoke first, out of
the corner of his mouth, slanting an insolent look up at the
ranchman.
“Youse de traffic cop on dis block, mister?”
Lon Forbes answered. “We know your sort an’ don’t want ’em here.
Shack! Hit the trail pronto! No back talk about it either.”
Cig looked at the big foreman. “Gawd!” he jeered. “Wotcha know
about that? De king o’ Prooshia on de job again.”
The bluff tanned Westerner took a step or two toward the ferret-
faced man from the slums. Hurriedly York spoke up. He did not want
anything “started.” There were stories current on the road of what
ranchmen had done to hoboes who had made trouble. He knew of
one who had insulted a woman and had been roped and dragged at
a horse’s heels till half dead.
“We ain’t doin’ no harm, boss. But we’ll beat it ’f you say so. Gotta
roll up our war bags.”
Reed did not discuss the question of the harm they were doing. He
knew that a spark might ignite the wheat, but he did not care to plant
the suggestion in their minds. “Put out the fire and move on,” he said
harshly.
“De king o’ Prooshia an’ de clown prince,” Cig retorted with a lift of
his lip.
But he shuffled forward and began to kick dirt over the fire with the
toe of his shoe.
Reed turned to the youngest tramp. “Get water in that can,” he
ordered.
“I don’ know about that.” Up till now the tramp called Tug had not
said a word. “I’m not your slave. Get water yourself if you want to.
Able-bodied, ain’t you?”
The rancher looked steadily at him, and the longer he looked, the
less he liked what he saw. A stiff beard bristled on the sullen face of
the tramp. He was ragged and disreputable from head to heel. In the
dogged eyes, in straddling legs, in the half-clenched fist resting on
one hip, Reed read defiance. The gorge of the Westerner rose. The
country was calling for men to get in its harvests. His own crops
were ripe and he was short of hands. Yet this husky young fellow
was a loafer. He probably would not do a day’s work if it were offered
him. He was a parasite, the kind of ne’er-do-well who declines to
saw wood for a breakfast, metaphorically speaking.
“Don’t talk back to me. Do as I say. Then get out of here.”
Reed did not lift his voice. It was not necessary. As he stood on the
bank above the sand bed he conveyed an impression of strength in
every line of his solid body. Even the corduroy trousers he wore
folded into the short laced boots seemed to have fallen into wrinkles
that expressed power. Close to fifty, the sap of virile energy still
flowed in his veins.
The fist on Tug’s hip clenched. He flushed angrily. “Kind of a local
God Almighty on tin wheels,” he said with a sneer.
York was rolling up his pack. Cig, grumbling, had begun to gather his
belongings. But the youngest tramp gave no evidence of an intention
to leave. Nor did he make a move to get water to put out the still
smoldering fire.
The rancher came down from the bank. Forbes was at his elbow.
The foreman knew the signs of old. Reed was angry. Naturally
imperious, he did not allow any discussion when clearly within his
rights. He would not waste his force on such a spineless creature as
York, but the youngest tramp was of a different sort. He needed a
lesson, and Lon judged he was about to get one.
“Hear me? Get water and douse that fire,” the ranchman said.
His steel-gray eyes were fastened to those of Tug. The tramp faced
him steadily. Forbes had a momentary surprise. This young fellow
with the pallid dead skin looked as though he would not ask for
anything better than a fight.

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Aquatic Animal Nutrition A Mechanistic

Perspective from Individuals to

Aquatic Animal Nutrition: Organic Macro- and Micro-

Animal Nutrition From Theory to Practice Philip Ian

An Introduction to Human Animal Relationships A

Animal Nutrition 8th Edition P. Mcdonald

Animal Cruelty Investigations A Collaborative Approach

Pathways to Our Sustainable Future A Global Perspective

The Human Evolutionary Transition: From Animal

Buddhism from a Catholic Perspective Paul M. Williams

Aquatic Animal Nutrition

Humboldt University at Berlin

ISBN 978-3-319-91766-5 ISBN 978-3-319-91767-2 (eBook)

Library of Congress Control Number: 2018953153

© Springer Nature Switzerland AG 2018

Berlin, Germany Christian E. W. Steinberg

1 Introduction – ‘You Are What You Eat’................................................... 1

3.2 Fishes................................................................................................ 80

3.2.1 Microbiome Ontogenesis...................................................... 81

6 Transgenerational Effects – ‘Your Offspring Will Become

6.1 Parental Effects................................................................................. 335

AAs amino acids

Autocrine (signaling) a form of cell signaling in which a cell secretes a signaling

chemical that binds to receptors on that same cell

Chaperon protein stabilizing new proteins to ensure correct folding

or helping to refold proteins that were damaged by cell

dbt encodes DOUBLETIME protein; a kinase that phosphory-

lates PER protein that regulates the molecularly driven,

fMHC fast myosin heavy chain, ATP-dependent motor protein,

involved in muscle contraction and other motility pro-

GRP gastrin-releasing peptide: a neuropeptide, stimulating the

Lecithotrophy nourishment and development of the embryo only via the

yolk originally contained within its egg; opposite:

Myogenin (= myog, myogenic factor 4) transcription factor involved

in the coordination of skeletal muscle development or

tim the negative loop of the molecular clock

RNAi RNA interference or RNA silencing: biological process in

which RNA molecules inhibit gene expression or transla-

tim a, b, h encode timeless proteins; essential in regulating the circa-

dian rhythm; part of a transcription-translation negative

Introduction – ‘You Are What You Eat’

‘You are what you eat!’

© Springer Nature Switzerland AG 2018 1

(Fig. 1.1) (Jordan 1905).

Fig. 1.2 A fly-eating frog

even spanning through evolutionary time.

hypothesis will surely challenge the paradigm of the so-called anti-nutritional

factors in animal feed in general, and for aquafeed in particular;

reducing potential constraints of production, and keeping it as productive, efficient,

of speciation. Alternatively, the action of microRNAs and other epigenetic

processes are beginning to be taken into consideration. In short-lived animals, such

Diets and Digestive Tracts – ‘Your Food

Abstract This chapter is an inventory of basic digestive mechanisms and of how

2.1 Digestive Tract

To understand the short-term, as well as the long-term and even transgenerational

© Springer Nature Switzerland AG 2018 9

incorporated numerous adaptations for the efficient breakdown and absorption of

Functional trophic group

by consuming a greater proportion of autochthonous material (algae and aquatic

macrophytes) instead of allochthonous material (terrestrial plant fragments and ter-

2.2.1 Protein Digestion

2.2.2 Lipid Digestion

Efficient lipid digestion requires emulsifiers in the mixture of food—mainly pro-

Berlin, Germany Christian E. W. Steinberg

1 Introduction – ‘You Are What You Eat’................................................... 1

3.2 Fishes................................................................................................ 80

3.2.1 Microbiome Ontogenesis...................................................... 81

6 Transgenerational Effects – ‘Your Offspring Will Become

6.1 Parental Effects................................................................................. 335

2.1 Digestive Tract

2.2.1 Protein Digestion

2.2.2 Lipid Digestion

2.2.3 Carbohydrate Digestion

2.3 Ontogenesis and the Intestine