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 Šíma1 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, Vídeňská 1083, 142 20 Prague 4, The Czech Republic
Corresponding author: Vaclav Vetvicka
Article Citation: Petr Šíma 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 reproduction in all medium, provided the original work is properly cited.
001-004 | JRB | 2015 | Vol 5 | No 1
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Šíma 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 Syvanen´s 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
Šíma 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
Ĺ Ăma 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 1622
Journal of Research in Biology (2014) 5(1): 1619-1626
Šíma 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 Peyer´s 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 Peyer´s 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: Peyer´s 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 (Šíma and Slípka, 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
Šíma 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
Peyer´s
some
patches.
functional Journal
of
into nuclear genome apparatus. These originally infectious Immunology. 125(6): 2626-2632. 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
Beltran P, Musser JM, Helmuth R, Farmer JJ, Frerichs WM, Wachsmuth IK, Ferris, McWhorter AC, Wells JG, Cravioto A and Selander RK. 1988. Toward a population genetic analysis of Salmonella: genetic diversity and relationships among strains of serotypes S. choleraesuis, S. derby, S. dublin, S. enteritidis, S. heidelberg, S. infantis, S. newport, and S. typhimurium. Proceedings of the National Academy of Science USA. 85(20): 7753-7757.
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