Journal of Research in Biology

ISSN No: Print: 2231 6280; Online: 2231- 6299

An International Scientific Research Journal

REVIEW

Journal of Research in Biology

Horizontal exchange of biological information and genesis

of adaptive immune system
Authors:
Petr ma1 and Vaclav
Vetvicka2

Institution:
1.University of Louisville,
Department of Pathology,
Louisville, KY 40202, USA

ABSTRACT:

Multicellular organisms have never evolved alone. Coevolution has been

accomplished by means of horizontal information transition. A wide row of
hierarchized interspecies interactions simultaneously transmitting biological
information had evolved in the past. In the present review, we summarize the current
hypothesis about possible horizontal exchange of biological information and genesis
of adaptive immune system.

2.Department of
Immunology and
Gnotobiology,
Keywords:
Institute of Microbiology,
Evolution, Genes, Bacteria, Vertebrates
Czech Academy of Sciences,
Vdesk 1083, 142 20
Prague 4, The Czech
Republic

Corresponding author:
Vaclav Vetvicka

Article Citation:
Petr ma and Vaclav Vetvicka.
Horizontal exchange of biological information and genesis of adaptive immune system
Journal of Research in Biology (2015) 5(1): 1619-1626

Web Address:
http://jresearchbiology.com/
documents/RA0481.pdf

Journal of Research in Biology

An International
Scientific Research Journal

Dates:
Received: 16 Sep 2014

Accepted: 19 Dec 2014

Published: 29 Jan 2015

This article is governed by the Creative Commons Attribution License (http://creativecommons.org/

licenses/by/4.0), which gives permission for unrestricted use, non-commercial, distribution and
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001-004 | JRB | 2015 | Vol 5 | No 1

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ma and Vetvicka, 2015

Horizontal exchange of biological information has a horizontal

information

transition.

Acquisition

of

horizontally transmitted genes explains the comments

substantial role in evolution

The first experimental evidence of the ability of written as early as 1970 (Anderson, 1970). This could be
microorganisms to incorporate the foreign genetic the answer to Syvanens query Why is molecular biology
material into their genomes was referred in 1944 (Avery so unified? (Syvanen, 1994).
et al., 1944). However, the idea of symbiosis of two or Defense of self vs. biological information exchange
more unrelated organisms, even pro- and eukaryotic, and

The

evolution

of

living

organisms

was

consequent exchange of their genetic material through principally the hierarchized mutual coevolution of units,
horizontal pathway, is much older. In 1867, Schwendener thermodynamically open systems (Bertalanffy, 1950), so
was the first to consider the role of intrinsic coexistence of called

the

CITROENS

(Complex

Information

two unrelated organisms in evolution. In 1879, De Bary Transforming, Reproducing Objects that Evolved by
introduced the concept of symbiosis, which was further Natural Selection), cooperating or competing with each
completed by Merezhkowsky into the Theory of other for space, nutrition, and survival (Orgel, 1973).
symbiogenesis in 1920. Then, since the end of the 20 th Their life and adaptive radiation were constricted by the
century, owing to the comparative work of Margulis horizontal exchange of biological information, and
(1993), it has been clear that the genes of mitochondria, simultaneously, by the defense of their internal integrity.
chloroplasts, and other organelles could be found Pathways of biological information exchange
incorporated in the genome within the nucleus of their
eukaryotic cells.

A
interactions

wide

row

of

hierarchized

simultaneously

interspecies

transmitting

biological

Multicellular organisms have never evolved information had evolved in the past, which by means of
alone, as evolution does not know a splendid isolation various molecules as information vectors and molecular
from other living beings. Not evolution, but coevolution mechanisms

in

formatted all organisms since the dawn of life. communication

the

role

among

of

receivers

non-related

allowed

micro-

and

Coevolution has been accomplished by means of macroorganisms (Table 1).

Table 1. Interspecific relations and interactions among organisms (The symbiosis sensu De Bary (1869))*
Interrelationship
Neutralism
Protocooperation

Mutual attitudes

Vectors

without mutual influence

none

relatively free, independent reciprocal profit

Attractants, allomones, kairomones, sinomones

Mutualism

fully dependent reciprocal profit

Attractants, allomones, kairomones, sinomones

Commensalism

relatively free profit for only one

Attractants, pheromones, kairomones

Amensalism
Antibiosis
Allelopathy

negative influence annihilation of an

organism defense to competition

Inhibitors, antibiotics, deterrents, phytoncides, repellents

toxins

Competition

mutual negative influence

Inhibitors, antibiotics, deterrents, phytoncides, repellents

toxins
Inhibitors, antibiotics, deterrents, phytoncides, repellents
Toxins, pathogenicity islands

Parasitism

attack toward an organism,

pathogenesis, gradual virulence

*) relations without definition of differences between parasitism and profit

1620

Journal of Research in Biology (2015) 5(1): 1619-1626

ma and Vetvicka, 2015

At least two pathways of biological information discharge enterotoxins increasing lesions (Butzler and
transfer could be regarded as fundamental:

1) the Skirrow 1979; Levine et al., 1983).

indirect pathway represents a number of factors and

Salmonella and Yersinia represent the third

signal molecules of various chemical compositions, group. These bacteria are able to translocate into intestinal
which are released into the environment (Table 1 and 2) mucosa, and proliferate in the lamina propria and
the direct instructive pathway of biological information mesenteric lymph nodes. Translocation of these bacteria,
transfer is represented by gene transmission, which is particularly S. typhi and S. paratyphi A and B could be
mediated by an amalgamation of entire genomes of two followed by systemic bacteremia (Levine et al., 1983;
or more organisms, insertion of smaller genome regions Beltran et al., 1988).
like single genes, and parts of genes (nucleotides). In this

From the point of evolutionary view, when these

case, the vectors represent mobile elements such as (and other) bacterial species capable of intracellular life
viruses (bacteriophages), conjugative plasmids, free overcome the host immunity, they could represent
nucleic acids (insertion elements, gene cassettes), potential vectors or direct donors of genes like
transposons, genomic islands, and possibly even prions Pathogenicity Islands (PAI), the groups of virulence genes
(Lee, 1996; Hacker and Carniel, 2001; Schmidt and on bacterial chromosome that could transfer new
Hensel, 2004; Prusiner, 1994; Silvestri and Baldassarre, biological information into the eukaryotic genome. The
2000).

PAI, the term introduced in 1990 by Hacker, may

Genomes are not protected by an immune system

represent mobile distinct molecular elements that take

It is not easy for a microbe, virus, or multicellular advantage of phage integration mechanisms. A question
parasite to enter into the internal milieu of a multicellular remains - represent bacteria like Salmonella, Francisella,
eukaryotic organism. The mucus and cell membranes of and similar intracellular microbes bearing PAI or other
the external and internal epithelia are effective barriers mobile elements, the contemporary evolutionary vectors
against the foreign invaders. In cases where these borders of new biological information? Could the infections
have been damaged, or invading parasites developed caused by these microorganisms in the instance that the
mechanisms to penetrate, a machinery of defense is victims survive have evolutionary consequences?
prepared

to

their

neutralization

and

annihilation. Advent of jaw vertebrates

Immunocompetent cells producing a row of defense

One of the great events of revolutions in the

molecules, including antibodies from immunoglobulin history of vertebrates was the appearance of jaws. The
superfamily, destroy the entire alien from outside.

importance of this evolutionary development can be

Generally, there are three groups of enteric hardly overestimated, for it opened new lines of
bacterial pathogens categorized on the basis of the adaptation

and

mechanisms by which they cause disease. The bacteria of advancement

to

new
the

possibilities
vertebrates

for
that

evolutionary
expanded

the first group, exemplified by Vibrio cholerae and immeasurably the potentialities of these animals. The
Escherichia coli, could be characterized by mucosal appearance of the jaws in vertebrates was brought about
adherence and production of enterotoxins, which cause by a transformation of anatomical elements that originally
diarrhea (Levine et al., 1983).

had performed a function quite different from the function

The bacteria of the second group (such as of food gathering (Colbert, 1980).
Shigella) penetrate and proliferate within the intestinal

Similarly, the same evolutionary importance

epithelia, which causes cell death. In some instances, they could be attributed to the emergence of the adaptive
Journal of Research in Biology (2015) 5(1): 1619-1626

1621

ma and Vetvicka, 2015

immune system. System in which not only ancient cells Activation Genes) allowing V(D)J recombination of
(like wandering and sessile macrophages), but also the molecules of immunoglobulin superfamily (Sakano et al.,
novelty specific only for deuterostomes, have started new 1979; Agrawal et al., 1998) (Table 3).
collaboration and cooperation utilizing and synthesizing Why RAGs transposed just into lymphoid cells?
new regulative and defense molecules in newly fabricated
structures, tissues and organs (Table 2).

The capability of translocation of intestinal

bacteria throughout the mucous lining and across

RAG time and emergence of the adaptive epithelia, and to penetrate into and survive within the
certain

immune system

developmental

stages

of

ancestral

rapidly

Some 500 million years ago, the predecessors of renewing cell populations like lymphoid cell lineage,
gnathostomes, the placoderms, emerged in primordial seas could represent mechanisms of a new genetic information
(Flajnik and Masahara, 2010). Possibly in these transfer in the form of gene cassettes like RAG
forerunners, the genomes of their immunocompetent cells transposons (Agrawal et al., 1998;Hiom et al., 1998;
were waiting for acceptance of new biological information Hansen and McBlane, 2000). On the other hand, the same
enabling them by means of highly sophisticated is
true
about
the
disability
of
primitive
mechanism of molecular recombination to transform immunorecognition of their hosts and a slightly effective
immunoglobulin superfamily molecules into effective immune response to destroy them.
defense vectors, the T and B receptors, antibodies, MHC,

It has been generally accepted that lymphoid

and other molecules needed for functions of the adaptive cells in a role of intraepithelial immunocompetent cells in
immune system.
all walls of the body make up at least a half of all the
Three events influenced the emergence of lymphocytes the mammalian immune system possesses.
adaptive immunity: evolution of gnathostomean basic They are loners slowly wandering free cells among the
body pattern inclusive jaws, differentiation of new epithelial cells. Evolutionary pressures like the threat of
immunocompetent structures inclusive lymphoid tissue decay of integrity of organisms caused by pathogens or
(particularly GALT), and acceptance of mobile parasites and to counter the effects of somatic potential
transposons of bacterial origin, the RAGs (Recombination carcinogenic mutations may be sufficient stimuli to
Table 2. Evolutionary consequences of gnathostomean basic body plan. Morphofunctional
prerequisites for adaptive immunity
Taxon

Cells

Organs

Molecules

Echinoderms

lymphoid-like

axial organ Ig

Protochordates

lymphoid-like

Ig

Chordates

lymphoid-like

stolon

Agnathans

lymphoid

typhlosole Ig

Gene cassettes incorporation into lymphoid cell lineage

Gnathostomes

lymphocytes

thymus

RAG, TCR, BCR etc.

GALT
bursa Fabricii-like
spleen
lymph nodes
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Journal of Research in Biology (2014) 5(1): 1619-1626

ma and Vetvicka, 2015

Table 3. Characters of Recombination Activation Genes (RAG1 and RAG2) suggesting horizontal transfer
- Unrelated in sequence, tightly linked
- Cleavage pathway similar to steps involved in DNA transposition reactions
- Retroviral type of integration
- DNA breakage and rejoining by means of trans esterification of mobile elements
- RAG and HIV integrases use alcohol instead of water as the nucleophile in the DNA cleavage reaction
develop rapidly renewing and specialized cell lineage that

In osteichthyan fish, the presence of gut

could neutralize these threats (Millar and Ratcliffe, 1989). lymphoid tissue has been documented, although they do
The first lymphoid-like cell populations endowed not possess structures like follicles resembling Peyers
with the primitive T and B diversity were documented in patches (Hart et al., 1988).

the axial organ of deuterostomian echinoderms, which is,

In amphibians, the first vertebrates, which have

moreover, supposed as phylogenetic homolog of the adapted to terrestrial life, the intraepithelial lymphoid
cells within the gut lamina propria and encapsulated

vertebrate spleen (Leclerc et al., 1993).

Urochordates contain several morphologically lymphoid structures in the oesophagus have been
distinct free cell types, but from the point of view of their described (Ardavin et al., 1982; Wong, 1972).
Intraintestinal epithelium of reptiles contains

morphology and immune functions, the most important

cell type is considered to be the lymphocyte-like cell, numerous

intraepithelial

lymphoid

cells

including

known also by name, the lymphocyte. The biological plasmacytes. Lymphoid cell aggregates in the gut
characteristics of these cells include the participation in submucosa resemble Peyers patches precursors in
encapsulation,

allograft

rejection,

cytotoxicity, mammals. Similar accumulations were found on body

proliferative response and immune memory (Sawada et locations of avian bursa of Fabricius or mammalian
al., 1993).

appendix (Solas and Zapata, 1980, Zapata and Solas,

In cyclostomes, the lymphohemopoietic cell 1979).

Birds possess the well-developed GALT exerting

aggregations accompanying the veins in the intestine

submucosa are considered as an early form of GALT. In many similarities to that of mammals: Peyers patches
these animals, which lack true thymus and spleen, these (Basslinger, 1858; Befus et al., 1980), cecal lymphoid
aggregations form about one-tenth of the gut wall volume nodules and tonsils (del Cacho et al., 1993; Glick et al.,
(Grozdinski, 1926, Tanaka et al., 1981). In the larval stage 1981),

and

in addition,

the

cloacal

diverticulum

of the ammocoete, the main lymphohemopoietic organ specialized to B cell development and production, the
most often homologized to vertebrate spleen or bone bursa of Fabricius (Glick, 1978; Jeurissen et al., 1989)
marrow is a longitudinal infolding along the anterior (Table 2).
intestine, the typhlosole (ma and Slpka, 1995) (Table

What

2).

gnathostomian vertebrates?
In elasmobranches, the first living gnathostomean

happened

before

the

emergence

of

It may be hypothesized with a high probability

vertebrates, the Leydig organ represents the important that the lymphoid cell lineage of predecessors of
lymphomyeloid structure associated to the anterior gut gnathostomes was sufficiently plastic for acceptation of
(Zapata 1981; Zapata and Cooper 1990).

RAG gene cassettes and for incorporation of them into

suitable loci of chromosomes. According to current

Journal of Research in Biology (2014) 5(1): 1619-1626

1623

ma and Vetvicka, 2015

knowledge, the sources of RAGs had to be some species 1980. Gut-associated lymphoid tissue in the chicken. I.
of bacteria that could enter into lymphoid cells, adapt to Morphology,

ontogeny

intracellular life, and transport a part of their chromosome characteristics

of

and

Peyers

some

patches.

functional
Journal

of

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bacteria could be endowed by a low degree of virulence
and might be capable to co-exist within cells for many
millions of years in some form of symbiosis like
protocooperation, mutualism or commensalism (Table 1).
Several questions remain. One focuses on
contemporary intracellular parasites such as Salmonella,
Shigella, Francisella etc., serving as potential vectors of
new biological information. Can they be considered an
evolutionary novelty? Are extant birds and mammals
bearing intracellular parasites in their highly organized

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