Improve your eel-farm

by David Kushnirov, Ph. D.

Foreword
Scientific Dogmas
Understanding eels solves problems
Rearing tank design
Micrographs of eel gonads
References

Contact Info: david.kushnirov@gmail.com

Foreword

Size variability develops in size-matched fish or in post-hatched fry. Studies on

trout fry led Brown (1957) to conclude that the presence of larger fish has a
suppressive effect on smaller companions, through the direct impact of
aggression on growth rather than through competition for food. Consequently,
growth of stunted fish is accelerated when the "jumpers tail", the fastest growing
cohort of the tank-mates, is removed. Brown (1957) called this phenomenon a
size hierarchy effect. Growth variation is typical for many species of fish and
crustaceans grown communally.

In the case of the eel, unequal growth of individuals associated with the "grading
problem", - is of great economical importance: 30-35 percent of elvers are either
discarded because of slow growth or held over to the next growing season in the
hope that growth will ‘pick up’. This brought Britain’s Fisheries Department and
Eel Producers Association in 1980 to give priority to the study of factors
influencing the growth rates of cultured elvers (Wickins, 1983).

Was there any progress since then in the understanding growth factors
associated with the grading problem?

Scientific Dogmas

When in 1986 I became involved in the study of the grading problem, I

encountered a number of dogmas circulating within eel farmers and fish
biologists.

The first dogma

In 1983, U. Wiberg showed the absence of sex (heteromorphic) chromosomes in
the European eel (Anguilla anguilla L.) and suggested an environmental
(metagamic) sex determination in the eel. Much earlier the same suggestion was
inferred from the highly skewed sex ratios observed in natural habitats. Quite
surprisingly and contrary to the clear indications of non-genetic sex-determining
mechanism in the European eel, the common belief was that sex in eels is
determined by the genetic mechanism, similar to one in mammals, birds and
some fishes (e. g. salmonids, common and grass carp, tilapias).
After acquainting miself with L. Bertin, F.-W. Tesch and other reviews on eel
biology I became convinced that environmental sex determination in eels is quite
plausible hypothesis.

Actually, the overwhelming majority of fish species are sexually labile; even
those, possessing the X-Y mammalian type of sex chromosomes, demonstrate
variant modes of sex determination (Francis, 1992).

The second dogma was a corollary of a concept that animal species have two
sexes - males and females, and that in eels females are larger than males.
Consequently, "the smaller eels in rearing tanks should be males".

The third dogma was also based on mere formal logic: since migrating silver
females are much larger than those of males and since in eel culture the fastest
growing cohort always consists of females, then the growth rate is a matter of
sex (like in humans, cattle and chickens). Consequently, since physiological sex
in fish is not determined ultimately by the genetic factors (contrary to humans,
cattle, chickens and fruit fly), hormonal feminization of a growing stock could be a
magic solution of the grading problem.

Actually, the grading procedure routinely used by eel farmers to promote growth
of slow-growers rejects a priori the existence of genetic determinants of individual
growth rate in eels. Effects of size, culling and social history on growth of cultured
elvers suggest that growth in elvers may be governed by behavioral or/and
physiological responses to handling and changing social environments (Wickins,
1987). Nonetheless, studies on genetic variation of slow and fast growing elvers
by polyacrylamide gel electrophoresis (PAGE) or DNA fingerprinting designed for
the revelation of the "selfish gene" (Degani & Gallagher, 1995) are more
prestigious than "primitive and cheap" deduction of the truth.

Surprisingly some students of eel biology conclude that males rather than
females grow faster at the sizes where the gonads are not completely
differentiated (Holmgren & Mosegaard, 1996).

As it can be seen, the matter becomes too ambiguous when "only" two sexes are
kept in mind: who grows faster, males or females? - Probably neither...?

G. Colombo et al. (1984) revealed that testis-like gonad of the yellow eel is more
primitive, and possibly reversible, than the frilled organ of Mondini - a
previtellogenic ovary. A testis-like gonad differentiates completely at the
beginning of sexual maturation and the metamorphosis into the silver (migrating)
male-eels (this never happens spontaneously in indoor tanks). Thus, the rearing
stock comprised of eels larger then 20 cm (the minimal length at which an ovary
was found) consists of females and undifferentiated (sexless) yellow eels, which
can differentiate into either females or males. This means that there are no males
in an indoor eel culture! That's, probably, why it is too hard for us to "catch a
head" of those who are not predestined to be boys or girls by their parents from
the "very beginning".

The fourth dogma

F. – W. Tesch (1977) cites a few descriptions, interpreted by him as a tendency
for eels to cluster in natural habitats: a few eels may hide together under the
same stone during the day (Krause, in Tesch, ibid.; Fig.79, p.210) or several eels
squeeze themselves into the same tube due to “tactile tendencies” (Tesch, ibid.,
p. 209), irrespective of the availability of additional unoccupied tubes placed in an
aquarium. Clustering of eels in culture tanks during daylight near heating and air
pipes is also a common sight interpreted generally by the same argument or "as
being gregarious”.

Indeed, several eels squeeze themselves into the same tube at the very
beginning of being placed in an aquarium... however in the course of time this
disposition changes (Fig.1) indicating some other "tendencies" governing eel's
behavior.

UNDERSTANDING EELS HELPS TO SOLVE PROBLEMS

One of the main problems, both
practical and theoretical, is size variation
developing during growth in rearing
tanks. It is this problem which prompted
my doctoral research in which I
approached this problem by linking
biological characteristics such as social
organization, growth performance, and
sexual development, including sex
determination, and viewing them as an
integrated entity.

1. Consideration of interactions between

three motivational systems - fear,
hunger and territoriality in respect to
shelters of different types revealed the
Fig. 1. Development of dominance within a dominancy of territorial aggression over
group of eels: A. a few eels squeeze other behaviors and solitarity as a social
themselves into the shelter at the start of the organization in this fish species during
experiment; B. the single occupant, the the immature phase of its life cycle. A
biggest eel in a group, appropriates the poorly structured and constrained
shelter. The resident of the shelter projects its artificial environment (e.g., a rearing
head and opens its mouth, threatening the tank) results in the transformation of
approacher. solitarity into a compound characterized
by the establishment of a linear ‘pecking’-order associated with both an increase
in frequency of agonistic encounters and a shift to more damaging patterns of
aggressive behaviour, resulting in the deaths of smaller lower rank contestants.
Availability of natural shelters (e.g., gravel) has the opposite effect by balancing
the mutually inhibiting relationships between the motivational systems, which
results in peaceful coexistence.

2. Better understanding of the causes of the eels’ aggressive behavior facilitated

a design of a rearing tank (Fig. 2) in which development of size variation is
minimized due to the decrease in frequency of aggressive interactions. Minor
additional installations in the standard rearing tank, namely, shelter which mimics
natural hiding-places, illumination, and a remote timer to control the
synchronization of feeding and light regimes, avoided the necessity of frequent
grading.
 Fig. 2. TANK DESIGN

Regular "bare" Ultra-structured

tank tank

Growth variation - Homogeneous growth

culling
Healthy fish, curative
Diseases caused by effect
social stress
All-female large fish
Prevalence of small
male-like fish In-tank bio-filter

Dependence on external Efficient feeding

bio-filter
Easy management
Food vestige

Inconvenient
management

This innovative technology in eel culture solves not only the grading problem, but
also the attaining of a higher growth rate, healthier fish and better utilization of
resources. Since the size hierarchy effect can be neutralized by application of the
method, it opens new perspectives for more precise studies on various
endogenous and exogenous factors influencing growth in eels (Kushnirov and
Degani, 1991).

3. Measuring cortisol level by radioimmunoassay (RIA) in the blood of eels

revealed low levels (less than 22 nmol/l) in blood of isolated eels and high
variation in eels sampled from the communal tank (from less than 22 nmol/l to
344 nmol/l). A negative correlation between eel size and blood cortisol content is
assumed. This implies an adverse effect of the General Adaptation Syndrome
(GAS) regarding health and growth of eels.

4. Light microscopy was used for revision of eel gonad differentiation, important
for understanding sexual dimorphism, sex determination and sex control in the
European eel, and has been a basic tool in the sections devoted to these
subjects. For mapping gonad development during ontogeny, tissue was sampled
by biopsy from immature (yellow) eels with induction of sexual maturity by a
single injection of human chorionic gonadotropin (HCG). The present findings
support the contemporary view that a testis-like Sirski’s organ is of bi-potential
sexuality, the main cell type of which are primordial germ cells (PGCs), arranged
in rows and cysts. This assumes a differentiated type of gonadal development in
the European eel, rather than undifferentiated type of development, proposed
elsewhere.

5. For the first time, sexual size dimorphism was clearly established in yellow
European eels on the basis of the regression of weight on length. Regression
analysis showed that within the size range of 24-47cm total length and 22-201 g
weight, female eels are on average lighter than sexless individuals with
undifferentiated gonads. The phenomenon is due to the loss of accumulated
weight associated with ovarian development up to the stage of basophilic
oocytes, spontaneous both in situ and in cultivo, in eels of particular size and
social rank. The difference in the Condition Factor (CF) between sexless and
female eels permits sexing of about 83% accuracy; this was used as a
complementary method to sexing based on macro- and microscopic appearance
of the gonad (Kushnirov and Degani, 1995).

6. Absence of heterochromosomes and a skewed sex ratio in natural habitats,

assuming an environmental sex-determining mechanism in anguillid eels,
promoted the experimental study on sex-determining factors. It appears that the
position in the social hierarchy determines the definitive sex, namely, sexless
high-ranking individuals are predetermined to become females, while submissive
eels differentiate into males; to verify the definitive sex, gonads differentiation
was induced by HCG injection (Degani and Kushnirov, 1992; Kushnirov, 1994).

For testes development induced by HCG see Micrographs of eel gonads.

7. Feeding elvers 60 mg/kg 17ß-Estradiol is found to induce femaleness,

confirming the lability of sexual development in eels. However, the method is not
practical for eel culture, as growth is effected by factors other than sex, namely,
by the amount of social pressure to which they are exposed.

A new concept, outlined in the present study - agonism in a compound of

A. anguilla eels is both a growth- and sex-determining factor. A space-time
structured artificial environment, reflecting the eel’s adaptation to benthic
life and solitary social organization, enables controlling the above
biological characters.

MICROGRAPHS OF EEL GONADS

Testis-like Sirski organ

a) Gonad cross-section (x260)

of yellow eel, 41 cm, 124 g.

At low magnification vermicular

appearance of the lobed organ is
confused with a tubular structure of
differentiating testes.

b) Higher magnification
(x1040) from Fig. a.

Rows of PGCs (arrows) are separated

by the irregular connective tissue.
Arteriolae with erythrocytes (Ery) are
seen.
Ovary of yellow eel, 41 cm, 91 g (x460)
Previtellogenic
basophilic
oocytes. Gross
macroscopic
appearance -
frilled organ of
Mondini

TESTES DIFFERENTIATION

Cross section through a seminiferous tubule of HCG-treated silver eel:

37,5 cm, 89g, (x650)
Formation of tubular structure is accomplished. The enlarged tubule wall consists
of large cysts with spermatagonia and spermatocytes. Lumen of seminiferous
tubule filled with spermatids and sperm cells.
BV - blood vessel
MATURE TESTES Cross section through a seminiferous tubule of HCG-
treated silver eel: 40.5 cm, 97 g, (x650)
Gross macroscopic appearance - overlapping lobes. Lumens of the thinned
seminiferous tubules are filled with sickle-shaped sperm.

References
Bertin, L., 1956. Eels, a biological study. Cleaver-Hume Press, London, 192 pp.

Brown, M.E., 1957. Experimental studies of growth. In Physiology of Fishes (M.E.

Brown,ed.), Academic Press, N.Y., Vol. 1, pp. 599-675.

Colombo, G., Grandi, G. and Rossi, R. 1984. Gonad differentiation and body
growth in Anguilla anguilla L. J. Fish Biol. 24: 215-228.

Degani, G. and Gallagher, L., 1995. Growth and nutrition of eels. Chapter 9.
Genetic variations in eels growing at different rates, pp.100-105, Laser Pages
Publishing Ltd. Jerusalem, Israel.

Degani, G. & Kushnirov, D., 1992. Effects of 17ß-Estradiol and grouping on sex
determination of European eels. The Progressive Fish-Culturist 54: 88-91.

Francis, R. C., 1992. Sexual lability in teleosts: developmental factors. Q. Rev.

Biol. 67: 1-18.

Holmgren K. & Mosegaard, H., 1996. Implications of individual growth status and
the future sex of the European eel. J. Fish Biol., 49: 910-925.

Kushnirov, D. & Degani, G., 1991. Growth performance of European eel (Anguilla
anguilla) under controlled photocycle and shelter availability. Aquac. Eng.
10:219-226.

Kushnirov, D. & Degani, G., 1995. Sexual dimorphism in yellow European eels,
(Anguilla anguilla L.). Aquaculture research, 26: 409-414.
Kushnirov, D., 1994. Individual growth in relation to agonistic behaviour, sexual
development and metagamic sex determination in the European eel, Anguilla
anguilla L., in aquaculture. Ph.D. thesis. Hebrew Univ., Jerusalem.

Tesch, F.-W. 1977. The eel. Biology and management of anguillid eels.
Chapman and Hall, London.

Wiberg, U.H., 1983. Sex determination in the European eel (Anguilla anguilla, L.).
Cytogenet. Cell Genet. 36:589-598.

Wickins, J. F., 1983. Speed-up for slow eels. Fish Farm, 6(2):24-25.

Wickins, J. F., 1987. Effects of size, culling and social history on growth of
cultured elvers, Anguilla angulla (L.). J. Fish Biol., 31:71-82.