copyright Jon Wood

For ordering information contact:

Fishingbooksender,
Caerbannog, Sarn,
Powys SY16 4EX
www.fishingbooksender.com or
www.carpfishingscience.com
Telephone 05601 972040

ISBN 978-0-9567297-1-2

The right of Jon Wood to be identified as the Author of the Work has been asserted by him in
accordance with the Copyright, Designs and Patents Act 1988. All rights reserved. No part of this
publication may be reproduced, stored in any retrieval system, or transmitted in any form, or by any
means, electronic, mechanical, photocopying, recording or otherwise without prior permission of the
publishers and copyright holders. No re-binding or alterations to this book without written permission
from the publishers and copyright holders.

Contents
Page
Introduction

1.

Water

What is Water?
The Soup
The Water Basin
The Main Characteristics of Water

4
5
8
9

a)
b)
c)
d)
e)
f)
g)
h)
i)

9
11
14
17
20
22
25
27
29

2.

pH
Dissolved solids
Hardness and alkalinity
Dissolved gases
Metals
Suspended solids
Phosphorus
Ammonia, nitrite and nitrate
Temperature

Lake Formation and Characteristics

33

Background
General lake information
Formation types

33
33
34

a)
b)

34
37

Natural processes leading to lake formation

Artificially formed lakes

Changing lakes
Light and heat
Stratification
Oxygen
Nutrients
Sedimentation

38
38
39
40
40
41

3.

42

Sediments

Background
Sediment types

42
42

a)
b)
c)

42
43
43

Silt
Clay
Gravel

Deposition

43

Sediment profile and benthic health

44

4.

Natural Food and Food Categories

46

a)
b)
c)
d)
e)
f)
g)
h)

Concepts
General diet
Herbivory
Waterborne food
Animals
Other foods arriving in the aquatic environment
Abundance and diversity in the aquatic environment
Constituents and chemical composition of major food items

46
47
48
51
53
65
65
67

Food categories

77

a)

Proteins

77

i)
ii)
iii)
iv)
v)
vi)
vii)
viii)
ix)
x)

Classification
Structure
Properties
Chemical determination
Gross protein requirements
Amino acids
Essential and non-essential amino acids
Essential amino acids and protein quality
Quantitative requirements of amino acid
Supplementing the diet with amino acids

77
77
78
78
78
79
79
80
80
81

b)

Carbohydrates

81

i)
ii)
iii)
iv)
v)

Classification and chemistry

Carbohydrate metabolism in fish
Digestion, absorption and storage of carbohydrate
Other factors affecting metabolism
Energy transformation

82
83
83
84
84

c)

Lipids

85

i)
ii)
iii)

Factors affecting the fatty acid composition of fish

Body lipid composition and dietary lipid requirements
Essential fatty acid requirements of carp

86
87
88

d)

Vitamins

88

i)
ii)
iii)

Water-soluble vitamins
Fat-soluble vitamins
Vitamin requirements of carp

88
90
90

e)

Minerals

91

i)
ii)
iii)

Calcium and phosphorus

Magnesium
Other essential inorganic elements

92
93
93

f)

Energy

94

i)
ii)
iii)

Energy flow in animals

Energy loss
Energy sources

95
95
96

a)
b)
c)

Fats
Carbohydrates
Proteins

96
97
97

iv)

Energy requirements of fish

97

a)
b)
c)

Energy distribution in relation to feeding level

Maintenance energy
Energy cost of growth and factors that alter energy needs

98
98
98

g)

Specific studies regarding natural food and food groups in carp

99

5.

Climate, Water Movement and Aquatic Dispersion

101

Introduction
Horizontal water movement
Vertical water movement
The effect of climate and water movement on environmental conditions and carp behaviour
Principles of diffusion and dispersion in water
Oil and water-based dispersion
Optimizing attraction

101
101
103
104
106
107
108

6.

109

The Carp

Background

109

a)
b)
c)
d)
e)
f)
g)
h)
i)
j)

109
110
111
111
111
112
114
115
116
117

Origins
Classification
Diversity, earliest carp and distribution
Habitat and general behavior
Physical appearance
Physical changes due to domestication and culture
Reproduction
Lifespan
General food and feeding habits
Predation

Responses to environmental changes

117

a)
b)
c)
d)
e)

117
119
121
122
123

Temperature
Oxygen
pH
Salinity
Habitat use

Swimming and Maneuvering

Breathing

124
125

Osmoregulation
Senses

126
127

a)

Sight

128

i)
ii)

Theory of vision and sight in fishes

Studies and information specific to carp vision

128
132

b)

Hearing

135

i)
ii)
iii)

Theory of sound in water

Theory of hearing in fishes
Studies and information specific to carp hearing

135
136
137

c)
d)

Smell
Taste

139
140

Food Location and Identification

Feeding

143
145

a)
b)

Why carp feed

The feeding mechanism

145
145

i)
ii)
iii)
iv)
v)
vi)
vii)
viii)

The development of the feeding mechanism in fish

Fish-food interactions
Functional morphology of feeding in teleosts
Feeding structures
Food processing
Food intake
Feeding abilities and diet of the carp
Conclusions

145
147
148
151
152
154
160
164

c)

Digestion

165

i)
ii)
iii)
iv)
v)

The digestive process

Protein digestion
Carbohydrate digestion
Fat digestion
Specific information relative to carp digestion

165
166
166
166
167

Growth

169

i)

Specific information regarding growth in carp

171

7.

Bait

177

Development of carp baits

Principal bait types

177
179

a)

Standard baits

180

i)
ii)
iii)
iv)

Bread
Worms
Corn
Tiger nuts

180
180
181
182

v)
vi)
vii)
viii)
ix)

Hempseed
Luncheon meat
Molluscs
Insect larvae
Summary and comparison of nutrient contents of standard baits

183
186
186
187
188

b)

Elaborated baits

193

i)

Boilies

193

Basics
Boilie theory
Boilie improvements
Factors affecting boilie effectiveness

193
194
196
197

ii)
iii)

198
200

Pellets
Artificial baits

Palatability
Additives and attractants

201
204

i)
ii)
iii)
iv)
v)
vi)
vii)

204
205
206
208
208
209
209

L-Amino acids
Minamino
Betaine
Inosine
Corn steep liquor
Butyric acid
Robin Red

Bait density
Specific information and studies related to carp baits and preferences
i)
ii)

Orientation of carp to free amino acids

Composition and nutrient digestibility of commercial baits

210
211
211
211

Bait conclusions

215

8.

216

Diseases and Carp Care

The stress response

Infection and transmission

216
219

a)
b)

219
220

Natural protection
Disease symptoms

Principal carp diseases

221

a)
b)
c)
d)
e)
f)
g)

222
223
224
225
226
226
227

Spring viraemia of carp (SVC)

Fish pox
Koi herpes-virus (KHV)
Columnaris disease
Carp erythrodermatitis (CE)
Fungal diseases
Parasites

Carp care

228

a)
b)
c)
d)
e)
f)

Rigs
Landing
Unhooking and weighing
Mouth and body treatment
Handling
Other carp care tips

229
229
230
230
231
231

9.

Conclusions

232

a)
b)
c)
d)

The Carp
The Carps Environment
Carp Location
Food and Bait

232
235
236
238

Bibliography

242

6.

The Carp

When youre fishing for one individual fish, you have to have tunnel vision. You have to have it in your thoughts 24/7
Ritchie McDonald in Advanced Carp Fishing.

Background
a)

Origins

The carp Cyprinus carpio is a fish native to Europe which has been introduced to every part of the
world with the exception of northern Asia and the poles. The original common carp was that found
in the inland delta of the Danube river about 2000 years ago, and was torpedo-shaped and goldenyellow in colour. It had two pairs of barbels and a mesh-like scale pattern (Balon, 2004).
Although this fish was initially kept as an exploited captive, it was later maintained in large,
specially built ponds by the Romans in south-central Europe (verified by the discovery of common
carp remains in excavated settlements in the Danube delta area). As aquaculture became a
profitable branch of agriculture, efforts were made to farm the animals, and the culture systems
soon included spawning and growing ponds.

Figure 6.1 A wild common carp (a) and its feral form (b) from the Danube delta in 1900 (from Antipa, 1909).

Unintentional artificial selection took place, and between the twelfth and fourteenth centuries, the
streamlined, fully scaled carp had transformed into a deeper-bodied, partially scaled or scaleless
domesticated version. The appearance of coloured versions (aberrations) became the koi of
Japan, but the common carp was not, as is often thought, originally domesticated in China.
Cyprinus carpio was therefore the oldest domesticated fish, and for that reason, we have had a
long relationship with the animal. Although initially the carp was a food item, in later times, and as
more fish species have become readily available for the table, the importance of carp culture in
Western Europe has become less important because of declining demand, partly due to the
appearance of more desirable table fish such as trout and salmon through intensive farming (see
the figure below), and environmental constraints. However, fish production in ponds is still a major
form of aquaculture in Central/Eastern Europe, including the Russian Federation where most of the
production comes from low intensity or semi-intensive ponds. In Asia, the farming of carp
continues to surpass the total amount of farmed fish volume of intensively sea-farmed species
such as salmon and tuna.

Figure 6.2 Inland aquaculture production of rainbow trout and common carp by volume in Europe
(source FAO, in Vradi, 2001)).

b)

Classification

Within the animal kingdom (Animalia) and the phylum Chordata, Cyprinus carpio is in the subphylum Vertebrata (vertebrate), class Actinopterygii and order Cypriniformes. The cypriniformes
(family cyprinidae) is traditionally grouped with the Characiformes, Siluriformes and Gymnotiformes
to create the superorder Ostariophysi, since these groups have certain common features such as
being found predominantly in fresh water and the fact that they possess Weberian ossicles, an
anatomical structure originally made up of small pieces of bone formed from four or five of the first
vertebrae; the most anterior bony pair is in contact with the extension of the labyrinth and the
posterior with the swimbladder. The function is poorly understood but it is presumed that this
structure takes part in the transmission of vibrations from the swimbladder to the labyrinth and in
the perception of sound, which explains why the Ostariophysi have such a great capacity for
hearing.

Most cypriniformes have scales and teeth on the inferior pharyngeal bones which may be modified
in relation to the diet. Tribolodon is the only cyprinid genus which tolerates salt water, although
there are several species which move into brackish water but which return to fresh water to spawn.
All of the other cypriniformes live in continental waters and have a wide geographical range
(Billard, 1995).
c)

Diversity, Earliest Carp and Distribution

The diversity of cyprinid species is at its greatest in China and South-East Asia, but is lower in
Africa and North America, where there are nevertheless 36 genera and 280 species (118
belonging to a single genus Nototropsis).
Fossilized cyprinids go back to the Plaeocene era in Europe, the Eocene era in Asia and the
Oligocene era in North America. The cyprinids are dispersed to the east of the Wallace line,
reaching Borneo, but have not crossed the Strait of Makassar, between Borneo and Indonesia. The
carp originating from Western Asia is no longer represented in the wild form other than by the subspecies Cyprinus carpio haematopterus in Eastern Asia and Cyprinus carpio carpio in Eastern
Europe (Billard, 1995).

d)

Habitat and General Behaviour

Carp exploit large and small manmade and natural reservoirs, and pools in slow or fast moving
streams. They prefer larger, slower-moving bodies of water with soft sediments but they are
tolerant and hardy fish that thrive in a wide variety of aquatic habitats (Page & Burr, 1991; Froese &
Pauly, 2002).
The attributes of carp include high environmental tolerances such as with regards to temperature
(2 to 40.6C), salinity up to about 14 parts per thousand, pH from 5 to 10.5, and oxygen levels as
low as 7% saturation. These, as well as being omnivorous, with a high reproductive capacity, are
reasons why the fish is widely distributed and found in most types of freshwater habitat.
Carp can typically be found in small schools, although larger carp often lead a solitary existence
(Smith, 1991).
e)

Physical Appearance

Carp often grow 30 to 60 cm in length and weigh 0.5 to 4 kg (Tomelleri & Eberle 1990); it is not
uncommon for common carp to reach 15 to 20 kg (McCrimmon 1968). Males are usually
distinguished from females by the larger ventral fin. Carp are characterized by their deep body and

serrated dorsal spine (Nelson, 1984). The mouth is terminal on the adult and subterminal on the
young (Page & Burr, 1991). Color and proportions are extremely variable, but scales are always
large and thick. Three sub-species with slightly different scale patterns are recognized. C. carpio
communis (scale carp) has regular concentric scales, C. carpio specularis (mirror carp) has large
scales running along the side of the body in several rows with the rest of the body naked, and C.
carpio coiaceus (leather carp) with few or no scales on the back and a thick skin (McCrimmon,
1968).
Carp usually possess 48-50 pairs of chromosomes, but some species such as the common carp
and the crucian carp have sometimes more. Hybridization occurs frequently in cyprinids, resulting
partly from environmental changes, in particular changes in spawning habitat: these changes can
be natural or due to human activities.
Two key physical features of carp are that they possess bilateral symmetry (having body symmetry
such that the animal can be divided in one plane into two mirror-image halves. Animals with
bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends) and the
lack of sexual dimorphism (the two sexes look alike).

f)

Physical changes due to domestication and culture

Mainly as a result of the addition of supplementary food to the carps environment, initially in ponds
and later in tanks, the species has undergone a number of morphological changes which now
result in a wide range of carp types available to the angler (see Hughes & Crow, 1997).
When the wild carp was introduced into pond systems, it naturally started to change its torpedoshaped body into a deep, laterally compressed and hunchbacked body. Soon individuals appeared
that did not have a regular, geometrical arrangement of scales, but showed severe irregularities,
scale-reductions or even complete scalelessness. These variations soon became a foundation for
artificial selection. Ultimately, domesticated common carp are represented by a variety of forms,
such as the fully scaled carp, linear, mirror carp and leather (or naked) carps (Brylnska, 1986;
Gorda et al., 1995; Pokorn et al., 1995).
The domesticated carp, which is of major importance in modern stocked fisheries, not only
changed its external shape, scalation and colour, but also underwent internal and physiological
changes (Balon, 1974, 1995a,b; Baru et al. 2002). Briefly, the mouth gape, the first character that
Rudzisky (1961) and Steffens (1964) used for clear distinction between wild and domesticated
common carp, was much smaller in the wild than in the domesticated varieties. Even more
pronounced differences are encountered among the mouth-gape indices calculated by dividing ten
times the mouth gape in cm2 by the length of head in cm: 4.46 to 5.57 for the wild common carp
and 8.12 for domesticated fish (Steffens, 1964). Both Rudzisky (1961) and Steffens (1964)
considered the enlargement of the mouth in domesticates to be caused by diet change, and
possibly the result of artificial selection. Domesticated common carp selected to utilize

supplementary food, grew better in ponds when man-made food was added (Sibbling, 1988).
Studies on the wild common carp in ponds (Rudzisky, 1961; Leszczysky & Biniakowski, 1967)
proved that wild common carp progeny was better suited for stocking natural riverine habitats than
were domesticated varieties.
Rudzisky (1961) found and Steffens (1964) confirmed that the intestine of the wild common carp
was 15 to 25% shorter than that of the domesticated carp. The index calculated by dividing the
length of the intestine by standard length (Ls) in cm was 2.11 for wild common carp and 2.64 for the
domesticated fish. Longer intestines in domesticated fish are probably due to utilization of
vegetable food not normally consumed by wild common carp. Both authors also found, when
comparing the deeper body of domesticated common carp with the cylindrical one of wild common
carp, that the calculated muscle mass was the same in both, although it appears to be greater in
the domesticated fish. This means that the dressed mass of the domesticated fish is not larger in
spite of the different body proportions. Also, the wild common carp has both chambers of the
swimbladder of similar size, whereas the domesticated fish has the anterior chamber always much
larger and the posterior chamber much smaller. This proportion (%) as given by Steffens (1980) is
61:39 for the wild and 90:10 for the domesticated. This may be related to the relative greater mass
of the head in a domesticated carp.

Figure 6.3 The author with a stocked mirror showing the relatively larger head and body form and scale pattern
which are far removed from the original wild carp stereotype.

The greater strength of the wild common carp is supported by some of its physiological attributes
(Steffens, 1964). Wild common carp have 18 to 19% more erythrocytes (red blood cells) and
haemoglobin (the oxygen-carrying component of the cells) than domesticated fish, and the blood

sugar level is 16 to 26% higher in the former. Also, wild common carp have a much lower water
content in muscles and liver than domesticated fish, greater fat content in individual organs, more
glycogen (the polysaccharide of glucose which forms the primary short-term energy source in
animals) in the liver, and more vitamin A (a vitamin with importance in vision and bone growth) in
the eyes, intestine and liver. In addition the muscles of the wild common carp are better
vascularized (contain more blood vessels) and do not fatigue as quickly as do those of
domesticated common carp (Balon, 2004). The persistence of these traits in some carp varieties
may be the reason why they present the angler with better fights than others.
g)

Reproduction

Carp generally spawn in the spring and early summer depending upon the climate. They segregate
into groups in the shallows to spawn. Carp prefer shallow waters with dense macrophyte cover.
Males externally fertilize eggs, which the females scatter over macrophytes in a very active manner
and the eggs stick to the substrate upon which they are scattered. A typical female (about 45 cm)
may produce 300,000 eggs, with some estimates as high as one million over the breeding season.
Incubation is related to water temperature and has been documented as being three days at
temperatures of 25 to 32C. Fry average 5 to 5.5 mm in total length. Temperature, stocking
density, and availability of food influence individual growth and by the time the fish reach 8mm, the
yolk has disappeared and they begin to actively feed (Froese & Pauly, 2002; McCrimmon, 1968).
H

Carp usually reach sexual maturity when the testes (in males) and ovaries (in females) develop.
Sexual maturity can occur at a very early age in carp where waters are constantly warm, but more
often, it is reached when i) a male is around 2 years old, ii) a female is around 3 years old, iii) they
exceed 30 cm in length and iv) weigh around 400 grams. (Swee & McCrimmon, 1966). Females
facilitate attachment of fertilized eggs to the substrate. There is no further parental care, the
hatching of the eggs is rapid and the newly hatched carp grow very quickly.
Because ovaries are much larger organs than the testes, females are generally easier to spot, as
the stomachs of mature females are mostly plump, while males remain a sleeker 'torpedo' shape.
When males are ready for spawning, they develop breeding nodules on the head and pectoral fins,
principally along the bones of the fin rays. These breeding nodules appear as fine whitish raised
spots. The nodules appear in abundance on the pectoral fins in regular rows and are rough to the
touch. During breeding, the male nudges the female with his head and fins to encourage spawning
(Swee & McCrimmon 1966).

h)
0B

Lifespan

There are reports of carp living more than forty years (see table below). Other cyprinid species are
included for reference, with the (+) symbol signifying that the fish was still alive when the age was
assessed:
Table 6.1 Summary of cyprinid longevities according to the Max Planck Institute for Demographic Research
Scientific Name

Common Name Wild Captive

Abramis brama

Bream

Abramis brama

Bream

Abramis brama

Bream

Carassius auratus

Goldfish

Carassius auratus

Goldfish

Cyprinus carassius auratus

Goldfish

10.0

Leuciscus cephalus

Chub

11.0

Leuciscus orfus

Golden orfe (+)

Rutilus rutilus

Roach

17.0

Reference
Altman & Dittmer (1962), Flower (1935)
H
H

10.0

Altman & Dittmer (1962), Seemann (1961)

H
H

23.0

Beverton & Holt (1959)

30.0

41.0

Carlander (1969), Flower (1925),

Moyle (1976)
Bobick & Peffer (1993)
Flower (1925)
Altman & Dittmer (1962), Flower (1935)
H
H

14.25
12.0

Flower (1925)
Altman & Dittmer (1962), Flower (1935)
H
H

Scardinius erythrophthalmus Rudd

Cyprinius carpio

Carp

Cyprinus carpio

Common carp

Cyprinus carpio

Common carp

Cyprinus carpio

Carp

Cyprinus carpio

Common carp

Cyprinus carpio var.

Prussian carp

10.0

Altman & Dittmer (1962) Flower (1925)

H
H

38.0

Hinton (1962)
47.0

20.0

Flower (1935)
McCrimmon (1968)

6.0
38.0

Flower (1925)
Hinton (1962)

6.4

Flower (1925)

The two main methods for ageing fish are the counting of the rings on the scales or on the otoliths
(ear bone). Otoliths are part of fishs inner ear or vestibular apparatus, and reside in the cranial
cavity. They are composed of calcium carbonate and protein. They function as sound receptors
and are also used for balance and orientation.

There are 3 pairs of otoliths or ear stones in the inner ear of a carp. The largest pair of otoliths, the
sagittae, is routinely used for aging, since the rings (annuli) can be counted in the same way as
those on a cut tree trunk. So too can those of fish scales, which adopt a similar pattern.
1B

a)

b)

Figures 6.4 a,b Methods of ageing; a) otolith and b) scale reading of annuli

The reading of otoliths is preferred to scale reading when the fish species does not possess scales
(as is the case with a leather carp) or when the scales are particularly small, for example in the
case of some flatfish such as plaice.
2B

i)
3B

General Food and Feeding Habits

Carp are primarily selective benthic omnivores that specialize on invertebrates that live in the
sediments (Lammens & Hoogenboezem, 1991). Newly hatched carp initially feed on zooplankton;
specifically rotifers, copepods, and algae (McCrimmon, 1968). Young carp feed on a variety of
macroinvertebrates including chironomids, caddis flies, molluscs, ostracods, and crustaceans
(McCrimmon, 1968). Adult carp are known to eat a wide variety of organisms including, insects,
crustaceans, annelids, molluscs, fish eggs, fish remains, and plant tubers and seeds (McCrimmon
1968, Lammens & Hoogenboezem, 1991). Carp feed by sucking up mud from the bottom, ejecting
it, and then selectively consuming items while they are suspended (McCrimmon, 1968). The
feeding areas of carp are easily recognized in shallow water as depressions in the sediment (Cahn,
1929).

j)
4B

Predation

Predators on carp include large fish such as pike, birds such as herons and mammals such as
otters (Britton & Shepherd, 2005). The colouration of the fish helps to camouflage it against their
surroundings so it can stalk its prey or hide from predators.

Responses to environmental changes

The main environmental changes to which carp can be exposed are changes in temperature and
the concentration of dissolved gases which were described in chapters 1 and 5. The effects of
decomposition and the addition of quantities of rain water into a lake or river, may also cause
changes in pH, which has a direct effect on the species.
a)

Temperature

Temperature is the most important environmental factor affecting the activity of poikilothermic
(creatures whose internal temperatures vary, often matching the ambient temperature of their
immediate environment) animals such as fish. Membranes are the first targets affected by change
of temperature and their lipid (fat) components respond immediately to this challenge. Expectedly,
a decrease in temperature results in a decrease in motional freedom of constituent phospholipids
and vice-versa, with resulting alteration in membrane fluidity. In order to maintain structural and
functional integrity of these structures, poikilotherms must counteract this affect of temperature and
maintain proper fluidity in the new thermal environment. It can be hypothesized that organisms
unable to adjust membrane physicochemical properties to temperature are not able to survive in a
changing thermal environment.
Fish use several methods in order to adapt to temperature change by adjusting the composition of
membranes, and studies have shown that fish accumulate phospholipid molecules in a cold
environment in order to adapt to periods of low temperature (Farkas et al., 2001).
The metabolic rate of all ectothermic (creatures that control body temperature through external
means, such as the sun, or flowing air/water) organisms is strongly dependent on temperature, as
well as body size. This relationship tends, however, to be clearer in aquatic organisms, where the
large thermal mass of water buffers the rate of change of environmental temperature experienced
by organisms (Clarke & Johnston, 1999).
The oxygen consumption rate of fish is directly proportional to the temperature of the water, since a
fishs biological processes occur faster in warmer water. This means that the fish breathes faster, it
digests food faster and therefore requires more food than it would if in water of a lower
temperature.

The resting oxygen consumption of teleost (bony) fish has been found to vary significantly with
temperature. The estimate of the factorial increase in resting metabolism comparing a
representative tropical (30C) and polar (0C) teleost ranged from 5.9 to 6.2, which indicates that a
typical tropical fish consumes roughly six times as much oxygen as does a typical polar fish and
therefore must find six times as much food per unit time simply to fuel resting metabolism.
One of the most contentious aspects of the evolution of teleost fish to cold temperatures, has been
that of metabolic cold adaptation which was theorized by a Danish physiologist Krogh. He
predicted that fish which have evolved to live at very cold temperatures should have developed or
evolved some kind of compensation for the metabolic-rate depressing effect of temperature. His
first experiment into metabolic cold adaptation involved a single goldfish (Carassius auratus), a
species closely related to the carp, and which was exposed to a series of temperatures and its
oxygen consumption rate measured (Ege & Krogh 1914). The experiment produced a roughly
exponential decrease in respiration rate as the temperature was lowered, and this relationship
became known as Kroghs normal curve. Initial experiments seemed to confirm the theory, but
since then, a number of other trials have not been able to confirm whether metabolic cold
adaptation exists.
As far as differences in resting metabolic rates between different fishes, there are significant
variations within the major groups. However, resting metabolism has been shown to correlate
strongly with the activity of the species, with resting metabolic rate increasing from less active to
more active species (Morris & North, 1984). Relatively speaking, carp are not particularly active
species, and were shown to have lower metabolic rates and lower resting oxygen consumption
compared with Salmoniformes (salmon and trout), Perciformes (perch) and Gadiformes (cod-like
fish) (Clarke & Johnston, 1999).
Food intake regulation in fish is a complex interplay between endogenous and environmental
factors (Peter, 1979; Fletcher, 1984), with water temperature being viewed as one of the main
environmental factors influencing feeding and growth (Brett, 1979; Elliot, 1982). For example,
Rozin & Mayer (1961) demonstrated that feeding by goldfish is temperature dependent, with a
10C decrease in water temperature (from 25C to 15C) inducing a reduction in food intake.
Most natural water temperature changes are not abrupt, and carp are able to adapt to a wide range
of temperatures. However, the information available would suggest that temperature has an
important effect on both carp biological activity and behaviour, both of which should concern the
carp angler.

b)

Oxygen

We have already seen that there is an important relationship between temperature and dissolved
oxygen concentration, and for that reason this section is somewhat of a continuation of the
previous one.
Dissolved oxygen is an important concern for fishery owners particularly in summer, where high
temperature results in low oxygen concentrations, since the warmer water has a lower maximum
saturation level than cooler water. Also, in ice-covered lakes in winter, where the water has no
contact with the atmosphere because of the ice, and when the ice is covered by snow which
doesnt permit photosynthesis of aquatic plants beneath it, the oxygen can be gradually consumed
by the respiring fish and other animals, and as a result fall to very low levels.
It is known that carp react to low levels of dissolved oxygen by increasing their activity in search of
better oxygenated water (Schperclaus, 1990). In some lakes, this water is often found where the
tributary or inflow enters the lake, and on frozen ponds or lakes, it is often in this area to which the
carp move when oxygen levels become particularly low.
In Winter, the opposite of the formation of the thermocline in Summer occurs, and it is frequently
the case that the warmest water is found in the deepest areas, and shallow water may be warmest
close to the lake bed. When carp congregate in Winter, in an area which displays more attractive
environmental conditions, be that temperature, oxygen, or a mixture of both, then the movement of
the bulk of fish causes a mixing of the lower layers with those cooler layers closer to the ice. As a
result, the temperature of the higher layers increases and the ice in that area begins to melt.
Radio telemetry (tracking) has been used to monitor carp movement in winter as a method of
ascertaining the reaction of the carp to various stress factors, leading to a better understanding of
carp behaviour (Bauer & Schlott, 2006). Work has shown that in the absence of oxygenation, carp
prefer the areas closer to the inflow, due to the higher oxygen concentration to be found there.
Also, carp have been shown to move towards and remain in more oxygenated areas, despite the
presence in some cases of lower temperatures to be found there, for example when the
oxygenation of the water has caused a mixing of warmer lower layers with those closer to the ice,
resulting in a decrease in water temperature. This study was repeated the following year and the
same results were found, with carp initially preferring the deeper parts of the lake in December and
then moving towards the shallow part of the lake near the inflow at the end of January. The fish
returned to the deeper parts of the lake in February. The carp involved in the experiment were
weighed before and after overwintering, and were found to lose weight during those months (as
high as 10% of body weight).
In some commercial fisheries and aquaculture production units, in order to avoid problems which
may result from unfavourable environmental parameters, an effort is made to ensure that carp face
optimal conditions. For example, hypoxia (lack of oxygen) resulting from low dissolved oxygen, as
well as low water temperature or fluctuations of ambient temperature cause a whole-body stress
response (Vianen et al. 2001; Van den Burg et al. 2005). Metabolic pathways are affected as well

as the immune system and significant hormonal changes have been observed in C. carpio (Le
Morvan-Roger et al. 1995; Van Raaij et al. 1996; Zhou et al. 2000). Moreover, complex interactions
of environmental parameters may occur. For example, responses to hypoxia exhibit a temperaturedependant component in carp (Stecyk & Farrell, 2002).
However, environmental conditions in lakes are not easy to control if at all and especially for
the overwintering of carp, it is dissolved oxygen which is the most crucial parameter. Ice and snow
covering the pond prevent contact with air and photosynthesis, while degradation of organic matter
continues and results in decreasing dissolved oxygen. In winters with long periods of ice cover
and/or ponds with a high loading of phosphorus and high degradation, dissolved oxygen may drop
to a level dangerous for carp. Even though carp may withstand dissolved oxygen as low as 0.6-0.7
mg/L in winter (Haas & Menzel, 2003), they clearly try to avoid such situations, by enhanced
activity as it is known from other species of fish. In summer, if dissolved oxygen drops to 3.0 3.5
mg/L, the carp stop feeding and search for better oxygenated water (Schperclaus, 1990). As one
would expect, this is the same in winter.
In fact, prior to the oxygen crisis event in the experiment by Bauer & Schlott (2006), the carp mostly
roamed the deeper parts of the pond and avoided the shallow areas near the shore. This is
consistent for findings of carp overwintering undisturbed and under good environmental conditions
(Bauer & Schlott, 2004). The depletion of oxygen increased activity and carp moved to the shallow
areas near the tributary. This clearly demonstrated the intensive search of the carp for better
oxygenated water. As the oxygen supply improved, carp returned to the deeper parts of the pond.
In fact, dissolved oxygen was highly correlated (P<0.001) with the activity of carp tracked using
telemetry.
The observation of the carp clearly suggested that oxygen was probably the most important stress
factor during this study, influencing the activity of carp in the lake in winter. Water temperature,
which was found to have the greatest influence on carp overwintering under good environmental
conditions (Bauer & Schlott, 2004), seemed to have no influence on activity under low dissolved
oxygen or the influence was masked by the overwhelming effect of dissolved oxygen in the study.
Compared with oxygen and despite the potential of even minor temperature changes to alter
activity in carp (Cooke & Schreer, 2003), changes in ambient water temperature exhibit a relatively
mild stressor (Tanck et al., 2000). This explains why even when temperature is extremely low as
a consequence of aeration single carp were tracked near the source of the oxygenation,
accepting very low ambient temperature for better oxygenated water.
As long as the lake is not too cold (>4C) cyprinids can still feed satisfactorily (Huet, 1986).
Ziemiankowski & Cristea (1961) found that foraging is limited in winter but not completely
suspended. This corresponds with the findings during the same radio tracking study that intestines
of carp caught in November (4.5C) and December (4.4C) contained food. By contrast, the empty
intestines of carp which had aggregated near the inflow in January (at the height of an oxygen
crisis) may be interpreted as, along with low temperature of 2.1C, a result of stress caused by low
dissolved oxygen leading to a cessation of feeding (Schperclaus, 1990).
The investigation of the potential influence of low dissolved oxygen and temperature on the activity
of overwintering carp using radio telemetry has not only given an insight into behaviour of stressed

fish in ponds and lakes, but has demonstrated the suitability of this technique for the evaluation of
stressors on the activity and behaviour of carp in their environment.
Beside this, other telemetry techniques have been used to evaluate the impact of stressors on
common carp. Long baseline acoustic telemetry has been used by Shin et al. (2003) to evaluate
the effect of dynamite explosions on the behaviour of common carp.

c)

pH

pH in the environment is a much less variable environmental parameter than dissolved oxygen and
temperature. Heavy rain can affect the pH of a shallow lake, and in some cases, nearness to the
inflow of a lake experiencing relatively large input of rainwater or water at least of a different pH,
could theoretically result in a pH gradient in a lake. However, studies on the avoidance of particular
areas of water due to the presence of a pH difference are few and far between, but the existing
information would suggest that carp may show preferences with respect to pH as they do with the
other water parameters described.
i)

Low pH

The physiological effects of acid water on fish have been studied (McDonald, 1983; Howells, 1984;
Wood, 1989). From these studies, it is clear that the key toxic mechanism of acid stress is
disturbance of electrolyte balance at the gills, and not internal acidosis (acidification of the blood).
Ultsch et al. (1980) found, in carp, that two instant step changes in water pH (from 7.4 to 5.1 and
from 5.1 to 4.0) resulted in declining plasma sodium and chloride concentrations and a progressive
reduction of arterial pH. A further decrease of the pH to 3.5 finally led to the death of the animals
within 24 hours. Other studies with various fish species have produced similar results (McDonald,
1983; Howells, 1984; Wood, 1989).
In a study by Van Dijk et al. (1993), carp were exposed to sublethal levels of water acidity, and
gradual acidification to pH 4.0 led to only minor transient changes in the measured blood
parameters and did not cause any mortality in the 2-day experimental period. Thus, carp are able
to survive water of pH 4.0 for 48 hours, without major physiological disturbances. In addition, in the
first month after their return to the holding tanks, there was no mortality among the experimental
fish.
Blood pH was shown to fall during periods of hypoxia and recover once the oxygen levels were
returned to normal in the study by Boeck et al. (2000b) into the effect of salt stress on the
resistance of common carp to low oxygen levels.

ii)

High pH

Natural alkaline waters are not as common as acidic ones. In the carps environment, however,
temporary high levels of pH can occur during hot summers as an effect of algal blooms, although
the effects have been studied in carp ponds, where pH has reached 10.0 or even more (Alabaster
& Lloyd, 1980).
The effect of high pH has been studied on carp larvae, using different pH levels between 7.8 and
10.3 (Korwin-Kossakowski, 1992) where growth was shown to be retarded in alkaline water, and
the results showed significant differences in growth rate for both length and weight and also
reduced survival of larvae at the highest pH.
These effects were attributed to pathological alterations of both anatomy and physiology, and also
affected feeding behaviour due to the water conditions, since anorexia had previously been
observed in alkaline waters by Witschi & Ziebell (1979).
Another possible reason for the slower growth is reduction of respiration, and Jezierska (1988)
reported that oxygen consumption of carp fry at pH 9.5 was 1.8 mg/O2 per hour compared with 0.7
mg/O2 at a pH of 10.3 (Korwin-Kossakowski, 1992). Respiration is reduced mainly by mucus
coagulation in the gills, which is a consequence of high pH levels (Daye & Garside, 1976, 1980;
Jezierska, 1988). Jezierska (1988) also found that a pH level of 10.0 was lethal to carp.

d)

Salinity

Carp are found in tidal areas of several European rivers including the coastal areas of the Northern
Caspian Sea, where large-scale attempts for extensive management of mirror carp in the brackish
waters of the Baltic coast have been made successfully (Kuliyez & Agayarova, 1984; Schildhauer
et al. 1992). Martemanov (1996) reported that carp can survive sodium levels of 176 mM (10.3
parts per thousand) for months and diluted seawater (0.3 to 3.0 parts per thousand) has been
reported to enhance the survival, growth and development of carp larvae (Lam & Sharma, 1985).
However, exposure of carp larvae to higher levels of diluted seawater (10 to 15 parts per thousand)
showed unfavourable effects with lower growth rates and increased mortality (Abo Hegab & Hanke,
1982, 1984; Schildhauer, 1983). Despite the fact that freshwater fish can occur in brackish water,
studies examining the physiological responses to increased salinity are performed mostly on
anadromous fish, while comparatively few studies have focused on stenohaline freshwater fish
such as the carp (Balment et al. 1987).
A study previous to that of De Boeck et al. (2000b) by the same authors (De Boeck et al., 2000a)
investigated whether the observed failures in dealing with higher salinities were due to an
increased energy demand which exceeded the available energy sources and internal energy
stores. Although some effects could be explained partly by the reduced food intake of the salt
exposed fish, a clear additional effect was caused by the salinity itself, because extra energy was

required to survive under these conditions. The study by De Boeck et al. (2000b) assessed if these
metabolic changes affect the resistance of common carp towards hypoxia and thus influence the
capacity of the carp to cope with any further environmental perturbations.
The capacity of fish to deal with reduced levels of oxygen in the environment differs considerably
among fish species. In general, cyprinids are more tolerant towards hypoxia than salmonids (Ott et
al., 1980; Ultsch et al., 1980). As with most fish, common carp are oxygen regulators, meaning that
they maintain their oxygen consumption at a constant level along a gradient of environmental
oxygen concentrations, until a critical oxygen concentration (Cc) is reached. Below this level,
oxygen consumption becomes proportional to the ambient oxygen concentration. The metabolic
responses to changes in oxygen availability vary, depending on the physiological state of the
animal, level of activity and temperature (Grieshaber et al., 1988; Burggren & Roberts, 1991).
Under conditions of stress, the Cc is likely to increase, reflecting the decreased capacity of the fish
to cope with environmental perturbations (Ultsch et al. 1980; De Boeck et al., 1995).
When challenged with decreasing environmental oxygen concentrations in their normal freshwater
environment, carp can regulate their oxygen consumption down to environmental oxygen
concentrations of 34 M. After 1 week in brackish water, carp are still able to maintain their Cc at
55 M but after 1 month of exposure, the Cc increases substantially to an oxygen concentration of
120 M.
In conclusion, studies have showed that the usual tolerance of common carp towards low
environmental oxygen concentrations is affected by long-term exposure to brackish water, and
critical oxygen concentrations shift to values that could occur in the natural habitat of common
carp. Ammonia excretion was also disturbed after long-term exposure to the brackish water.

e)

Habitat Usage

Some examples of environmental factors affecting carp movement in lakes have been shown by
way of radio telemetry. Some attempts have been unsuccessful (Okland et al. 2003), but the
majority have shown some interesting findings regarding carp preferences and the ways in which
they avoid stressful conditions.
Studies related to thermal discharges from power stations have indicated that carp show a
preference for warmer water and have been shown to reside there for long periods of time (Cooke
& McKinley, 1999). The potential effects of residing in a thermal discharge channel include altered
metabolic rates, feeding responses and reproductive potential, increased susceptibility to disease
and at worst, death from cold or heat shock (Coutant, 1970; Spigarelli et al. 1974).
During the winter of 1997, the behavioural response of fish exposed to fluctuating discharge
temperatures in the NTGS discharge canal (Lake Erie) was examined (Cooke & McKinley, 1999).
A fixed radiotelemetry system was used to continuously monitor movements of common carp, for
several months in order to evaluate the distance fish penetrated and the duration of their residence

in the discharge channel, and correlate activity and residency to environmental and operational
conditions.
As a result of the radio-tracking, it was established that the common carp were resident in the
discharge canal for the majority of the winter of 1997-1998. Compared with another species
(Channel Catfish, Ictalurus punctatus), another bottom living fish, residency times within the
warmer water were greater, therefore showing a selective preference for the discharge water, with
carp spending almost 50% of their time in there rather than in the cooler lake itself.
Another study by Yoder & Gammon (1976) showed a similar preference for carp, where they were
found during the winter in much larger numbers in the heated effluent and backwater areas of
another electric generating station, rather than in the ambient river reference sites.
At other times of the year, carp have also been shown to prefer the warmer discharge environment.
In one study (Romberg et al. 1974), carp were shown to appear in large shoals in May and
routinely swim in and out of the discharge flume through temperature gradients as much as 10C.
Although the carp seems to seek out more comfortable conditions, their presence in heated
waters is unusual since these flows are not of a constant temperature. The water may be of a more
attractive temperature than that of the lake, particularly in winter, and it appears that as long as the
fish are not subjected to thermal shock, where great differences in temperature occur after
acclimatizing to a particular temperature over a period of time, then they still seem to avoid
stressful conditions.
This same study by Cooke & McKinley (1999) suggested that carp were attracted to the discharge
canal during periods of higher flow, therefore indicating that the carp knew that those flows would
be associated with warmer and more advantageous living conditions. By the same reasoning, an
angler may take advantage of this, since the evidence suggests that not only temperature but also
oxygen and maybe additional environmental conditions affect where the carp move to at different
times of the year and times of the day.
This suggests to the angler that if carp are able to selectively search out, identify and then occupy
areas of a lake with more appropriate environmental conditions, then identifying the areas of a lake
which offer those conditions at a particular moment, will allow him to increase his or her chances of
success.
Swimming and Maneuvering
Fish move through the water using their fins for locomotion, stability or balance, and steering. The
tail, or caudal fin, helps to propel the fish forward as it is moved back and forth - the actual forward
thrust coming from the pressure of the fish's tail against the surrounding water. Fish with smaller
caudal fins undulate their bodies to move forward.