2019 SCIENCELINE

Journal of Life Science and Biomedicine

J Life Sci Biomed, 9 (5): 144-150, Sept 2019

License: CC BY 4.0

ISSN 2251-9939

DOI: https://dx.doi.org/10.36380/scil.2019.jlsb23

Systematic review on avian immune systems

Mastewal BIRHAN¹^

College of Veterinary Medicine and Animal Science, Department Veterinary Paraclinical Studies, University of Gondar, Ethiopia

Corresponding author’s Email: maste675@gmail.com ; ORCID: 0000-0002-0984-5582

ABSTRACT

Review Article

PII: S225199391900023-9

Aim. The aim of this review paper is too summarized and compares avian immune systems

to the other domestic animals as comparative immunology type of review. Appreciation of

the avian immune systems and their functions are very critical for disease diagnostics and

Rec. 06 June 2019

Rev. 25 August 2019

Pub. 25 September 2019

new vaccine developments. Some of the avian immune systems are differ from mammalian

immune systems, based on their production sources of immune cells like B-cells production

site bursa of fabrics, but in mammalian is bone marrow. When we see the antibody type of

birds; there are three principal classes of antibodies: IgM, IgG, IgY and IgA. Antibody

diversity is achieved by gene re-arrangement. The other effector immune cell of birds is T

cells. There are two distinct pathways that are α/β and γ/δ, avian T-cell diversity is probable

made through combinatorial and junctional mechanisms. Recently, genes of several avian

cytokines have been cloned and expressed. A number of naturally occurring viruses cause

immunosuppression in chickens. Conclusion. There is much current interest in

Keywords

Antibody,

Avian,

T cells,

understanding the mechanisms of immunosuppression and developing strategies to Vaccine

enhance immune responsiveness in commercial poultry.

INTRODUCTION

One of the wonderful rules in the poultry industry is to hardly working on disease and predator control, good

institutional linkage, and with good management from the healthy birds it is possible to increased high

productive efficiency, capacity and with it, economic profitability” [1 ]. Scientific research on poultry

immunology and the diseases affecting avian species is not a new concept [2 ]. But, more recently, the chicken

was the first agricultural species as an income sources for which indicted by a genome sequence map [3 ].

Meaningful what specific immune molecules are encoded in the chicken genome delivers an outstanding

background to form and magnify our information on the avian immune systems [2 ]. Comparable other avian

immune systems, the immune system of chickens is made up of two types of mechanisms non-specific and

specific [4 ]. The potential pathogen and other risks facing mechanisms are slight different from those come

across by mammals. It is therefore essential that mechanisms be available to combat invading bacterial, viral

and parasitic pathogens and to destroy neoplastic or other altered cells. It is also essential in birds, as in

mammals, that the resulting immune response be regulated to ensure that it is adequate in quantity and quality

[5 ].

We need to understand the chicken immune system, to familiarize you with those defense mechanisms.

The bursa of Fabricius and the thymus organs are the central lymphoid organs in the chicken, essential to the

development of adaptive immunity [6 ]. In bird’s poor of all bursal lymphoid tissue, but still holding a normal

thymus, no circulating antibody was detected after challenge with different antigens. Delayed hypersensitivity

reactions to tuberculin or vaccinia virus (VACV) were nearly completely inhibited [7 ].

From the pronounced important avian organs, gut-associated lymphoid tissue is one of the organ that

contains functionally immature T and B lymphocytes at hatch, and that function is achieved during the first 2

weeks of age as demonstrated by mRNA expression of both ChIL-2 and ChIFNγ confirmed by Bar-Shira et al. [8 ].

The gut is a vital organ system which makes up two equally important functions, that are digestion systems

and host defiance [9 ]. When we address the chickens immune systems, the innate immunity includes physical

barriers (skin, mucus coat of the GI tract), specific molecules (agglutinins, precipiacute phase proteins,

lysozyme), phagocytic function of phagocytes (macrophages and neutrophils), and lysing activity of a class of

lymphocytes called natural killer (NK) cells [10 ].

In females birds, may improve their reproductive victory by mediating brotherly competition and growth

of offspring by means of differential hormone transfer to the egg yolk [11] [12] and [13]. For example, differential

Citation: Birhan M. 2019. Systematic review on avian immune systems. J. Life Sci. Biomed. 9(5): 144-150; www.jlsb.science-line.com

144

transfer of steroids to eggs within the same grasp may alleviate or intensification the effect of hatching

asynchrony as yolk steroids enhance nestling growth and competition [14] and [13]. Yolk testosterone was also

present in the eggs of female canaries that were kept without a male, indicating that it is of maternal origin [11 ].

Birds are born with an imperfect immune system and young chicks have to rely on maternal antibodies and the

innate immune defiance system to fight off pathogens [15 ].

While the avian system shares several similarities with mammalian systems, there are differences in the

genes and molecules involved, the cells and organs involved, as well as the functional mechanisms. Chickens, for

example, have a different assortment of Toll-like receptors, defensins, chemokines and antibodies. Birds do not

have eosinophils though the functional corresponding to the mammalian neutrophil is the avian heterophil.

Birds do not have lymph nodes, but do have a Bursa of Fabricius, which mammals do not. The mechanisms by

which the different receptors are generated are also fundamentally different [16 ]. Therefore, the aim of this

review paper are compering and analyzing of how the avian immune systems, structures organization, cells and

organs differ from the other domestic animal immune systems.

OVERVIEW OF THE AVIAN IMMUNE SYSTEMS

Studies that comprehensive study type, as a comparative method to the immunology with an gratefulness for

physiological ecology and evolution are defining an important new field in biology-ecological immunology [17 ].

Though in wide-ranging terms the avian immune response is strangely similar to that of mammals, when one

looks at specifics birds have a different repertoire of immune organs, cells and molecules compared to those

characterized in mammals.

The unique structures of chickens are adversely distresses by heat stress, so, due to this impressions

reeducations of productive performance, immune response, survival and profitability of fast growing chickens

[18 ]. Beyond the beneficial features, the risk regarding the development of antimicrobial resistance and

transference of antibiotic resistance genes from animal to human microbiotaled the European Union to ban the

application of antibiotics as growth promoters since1^stJanuary 2006, which was followed by the other parts of

the world including North America [19 ].

Avian are extremely vulnerable to varies infection by opportunistic pathogens during the first few days

after hatching [20 ]. In avian species, adaptive immunity encompasses both humoral and cell-mediated immune

(CMI) responses [21 ]. The avian embryo provides numerous compensations for studies on development of the

immune system [22 ]. The bird egg is worthily adapted to house, feed, and protect the developing embryo. The

outer lime-flavored shell and adherent shell membranes provide a physical barrier that excludes most

microorganisms, but permits free exchange of respiratory gases [23 ]. Interior to the shell membranes is a thick

zone of albumen that provides a sterile fluid medium for the free growth and morphogenesis of the embryo and

the extra embryonic membranes. In the center of the egg is the yolk mass that will nourish the embryo through

the incubation period [24 ].

Commonly, Birds are lack organized lymph nodes, yet have the Bursa of Fabricius. Birds lack neutrophils

and functional eosinophils, yet have a distinct group of polymorph nuclear granulocytes known as heterophil.

Birds also have a different repertoire of cytokines, chemokines, Toll-like receptors, defensins and integrin’s [25 ].

INNATE IMMUNE SYSTEMS

Innate cells

The innate immune system develops in the bone marrow (BM) from common myeloid progenitors (CMPs).

Due to the expression of AR in hematopoietic progenitors, there is reason to believe testosteronemay play an

important role in shaping the immune cell repertoire even prior to the cells leaving the BM [26 ].

Macrophages

Macrophages instigate from bone marrow stem cells by differentiating into monoblasts, promonocytes,

and monocytes. However monocytes establish foremost phagocytic cellular component in chicken blood, tissue

macrophages are extensively dispersed and present in almost every organ. Monocyte cultures from peripheral

blood leukocytes can be established by incubating the leukocyte fraction on a solid substrate such as Petri

dishes or glass coverslips. The adherent blood monocyte cultures can then be established by incubation and

washing off the non-adherent cell fractions [27 ]. The two most commonly used avian macrophages cell lines are

Citation: Birhan M. 2019. Systematic review on avian immune systems. J. Life Sci. Biomed. 9(5): 144-150; www.jlsb.science-line.com

145

MQ-NCSU. and HD11, an avian myelocytomatosis virus (MC29) transformed chicken macrophage-like cell line

[28 ]. The MQ-NCSU cell line was established from spleen of a broiler-type chicken experimentally challenged

with the JM/102W strain of Marek’s disease virus. The cultural, morphological and functional characteristics of

the MQ-NCSU cell line imply that this is a malignant-transformed chicken cell line belonging to the

mononuclear phagocyte lineage. Avian macrophages harvest chemotactic cytokines of both macrophage

inflammatory protein (MIP) families. The chicken MIP-1 and MIP-2 chemokines have the identical amino acid

motifs as mammalian chemokines: adjacent cysteine’s (CC) in the MIP-1 chemokines and cysteine’s separated by

another amino acid (CXC) in the MIP-2 family. The chicken MIP-2 family chemokine is currently designated as

9E3/CEF4. It has high homology to mammalian interleukin (IL)-8 and is abundantly expressed by activated

peripheral blood monocytes ([29]; [30] and [31].

TLR

Pattern recognition receptors (PRRs) are a serious component of pathogen recognition in both mammals

and chickens [32 ]. Toll-like receptors (TLRs), a main family of PRRs, are expressed in chicken intestinal tissues

and the local immune cells have been shown to respond to bacterial ligands [33 ]. Influxes of heterophil as well as

increases in cytokines and chemokines are evident [34 ] and [35 ] and are thought to contribute to the pathology

detected. However, once chicks are more than a few days old, S. Typhimurium persistently colonizes their

intestines in the absence of pathology [36 ], signifying that maturity of host defenses contribute to the deficiency

of clinical signs

In chickens, it has been established that heterophil constitutively express TLR2A, TLR2B, TLR1/6/10 mRNA

and that heterophil isolated from neonatal chicks and exposed to LTA undergo an oxidative burst [37 ]. There are

also data to suggest that CD14 and TLR2 mediate LTA-stimulated oxidative burst in heterophil [37 ]. Chicken

TLR3 expression pattern appears to be similar to what is observed in mammals [33 ]. For example, chicken

heterophil express TLR3 and are approachable to poly I:C, demonstrated by an induced oxidative burst and

degranulation of the stimulated heterophil, which may be mediated by a signalling pathway involving

phospholipase C, phosphatidylinositol 3-kinase and intracellular Ca²⁺⁺[38 ]. In contrast, others have confirmed

the poor ability of poly I:C to stimulate nitric oxide (NO) production in chicken monocytes, while HD11cells, a

chicken macrophage cell line, were readily stimulated to produce NO by poly I:C [34 ] and [39 ].

The first groups of TLR are expressed on the cell surface and recognize primarily cell-surface PAMPs. They

include TLR1, TLR2, TLR4–6 and TLR10 in human and TLR11 in mice. There are direct chicken orthologous of

mammalian TLR4 and TLR5 [40 ] and [33 ]. In mammals, there is a single TLR2 gene, and the genes encoding

TLR1, 6 and 10 lie in a single locus. Mammalian TLR2 forms functional heterodimers with at least TLR1 and

TLR6, allowing recognition of a wider panel of pathogen associated molecular patterns (PAMPs). At the

equivalent locus to the mammalian TLR1, 6 and 10 locus, the chicken genome encodes only two genes, TLR1LA

and TLR1LB [41 ] and [42 ]. Avian TLR repertoire and the response to various agonists [43 ]. Toll-like receptors

(TLRs) are important for eliciting innate immunity in animals by playing an essential role as pattern

recognition receptors that detect infectious pathogens by recognizing the conserved molecular structures

known as pathogen associated molecular patterns [44 ]. There are ten avian toll-like receptors and that five of

these, TLR2a, 2b, 3, 4, 5 and 7, are clear orthologous to TLRs found in mammals [45 ]. The non-mammalian TLR21

exists in many species of birds, fishes, and frogs [46 ] and [47 ]. As a homologue of mammalian TLR9, TLR21 can

recognize synthetic oligo-deoxy-ribo-nucleotides (ODN) and DNA viruses that contain CpG motifs, which

further trigger the innate immune response [46 ] and [48 ]. The RLR family encompasses three members: RIG-I,

melanoma differentiation-associated gene 5 (MDA5) and research laboratory of genetics and physiology 2

(LGP2), which are located in the cytoplasm

AVIAN ADAPTIVE IMMUNITY

Cell mediated immunity and humeral immunity

Chicken αβ T cells express either CD4 or CD8 accessory molecules, whereas most of the γδ T cells do not

[49 ]. The cytotoxic T lymphocyte response can decrease viral shedding in mildly pathogenic avian influenza

viruses, but provides doubtful protection against HPAI. Influenza viruses can directly affect the immune

response of infected birds, and the role of the Mx gene, interferon’s, and other cytokines in protection from

disease remains unknown [50 ]. Avian T cell progress has emerged with the use of monoclonal and functional

antibodies to elucidate T cell differentiation antigens and molecular and functional explanations of mammalian

Citation: Birhan M. 2019. Systematic review on avian immune systems. J. Life Sci. Biomed. 9(5): 144-150; www.jlsb.science-line.com

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T cell receptors (TCRs) [51 ]. Avian T cells bearing a γδ TCR are the first to be generated during ontogeny and

they comprise up to 50% of the recirculating T-cell pool in mature birds [52 ].

Progress of B cells in chickens proceeds via a series of disconnected developmental phases that includes

the maturation of committed B cell progenitors in the specialized microenvironment of the bursa of Fabricius

[53 ]. Three classes of chicken immuno globulins have been identified immunochemically [54 ] and genetically

[55 ] as homologues to the mammalian IgM, IgA and IgG, and their organizational properties have been reviewed

in more detail elsewhere [56 ]. The intestine is a complex tissue that includes a major immune constituent.

Indeed, the numbers of immune cells found in intestinal tissues exceed the numbers found in the rest of the

body [57 ] and [58 ].

Expression of selected genes involved in pathogen detection and the innate immune response were

profiled in caecal tissues by quantitative RT- PCR. TLR4 and TLR21 gene expression was transiently increased

in response to both bacterial species [59 ] and [60 ]. Defense of the intestinal mucosal surface from enteric

pathogens is initially mediated by secretory IgA (SIgA) [61 ].

Three classes of chicken immunoglobulin’s have been identified immunochemically [62, 63 ]and genetically

[64 ] and [65 ] as homologues to the mammalian IgM, IgA and IgG and their structural properties have been

reviewed in more detail elsewhere [66 ]. Chicken IgM is structurally and functionally homologous to its

mammalian counterpart, being current in serum as a high molecular weight pentamer of m2L2 units and being

the first antibody generated during a primary antibody response. IgM is also the major class of immunoglobulin

expressed on the surface of chicken B lymphocytes [67 ].

Chemokines and Cytokines

Interferon’s (IFNs) are a family of multifunctional cytokines with significant roles in cellular resistance

against viral infection [68 ]. In response to virus invasion, host pattern recognition receptors (PRRs) detect

pathogen associated molecular patterns (PAMPs) and subsequent activation of innate immune system through

retinoic acid inducible gene I (RIG-I) like receptors (RLRs)-MAVS-dependent IFN signaling or toll-like receptors

(TLRs)-TRIF/MyD88-dependent IFN signaling [69 ], eventually, inducing the expression of type I IFNs. IFNs then

bind their cognate receptors, triggering a signaling cascade that outcomes in the expression of abundant

interferon-stimulated genes (ISGs) by the JAKSTAT signaling pathway, various of which possess antiviral

properties [70 ] and [71 ]. Interferon regulatory factors (IRFs), a family of transcription factors, play authoritative

roles in the regulation of IFN expression during viral infection [72 ]. To date, 9 IRF genes (IRF1-9) have been

described in mammals, a tenth (IRF10) is present in numerous avian species and a total of 11 IRFs (IRF1- 11) have

been identified in fish [68 ].

CONCLUSION AND RECOMMENDATIONS

The chicken, perhaps surprisingly, has made several influential contributions towards our understanding of

immune responses as comparable way. Notwithstanding this, before the chicken genome sequence, our ability

to study immune systems in detail in birds is appropriate and also in our thoughtful of the immune gene

catalog. There are still gaps, both in the chicken immuno systems and their catalog. In comprehensive study is

well important by comparing of birds immune systems with the other animals. Both innate and adaptive

immune responses, with the latter including both cell-mediated and humoral immune responses, leading to

address and increases our knowledge about chicken’s immune activity. However, looking at the organs, cells,

and molecules of the immune response in birds, it appears that mammals and birds accomplish the equivalent

overall responses-often in quite different ways. Instead, we concentrate on the basic immune response, as well

as a description of the major cell types and major areas where the cells and molecules of the immune response

differ from those of mammals. Generally, the immune system of the chicken is very helpful in avoiding disease

and helping to insure maxi-mum productive potential is realized. We must learn how to take advantage of all

parts of the system when designing health programs. Based on the above information’s the following

recommendation will be forwarded:

 Researcher should be focus on the avian immune systems and their role of contributions, and the

delineation of the bursal and thymus-derived arms of the immune system.

 The genes of several avian cytokines have been cloned and expressed; so that scientists would be given

an attention for new disease resistant gene formed.

 The researcher should be emphasis on current attentiveness in thoughtful the mechanisms of

immunosuppression and developing approaches to advance immune responsiveness in commercial poultry.

Citation: Birhan M. 2019. Systematic review on avian immune systems. J. Life Sci. Biomed. 9(5): 144-150; www.jlsb.science-line.com

147

DECLARATIONS

Acknowledgment

The authors’ heartfelt thanks to University of Gondar, research and community service v/president office,

college Veterinary Medicine and Animal sciences for the financial and resource supporting

Authors' contributions

Mastewal Birhan conceived the review, coordinated the overall activity, and write and submit the

manuscript.

Availability of data and materials

Data will be made available up on request of the primary author

Consent to publish

Not applicable.

Competing interests

The authors declare that they have no competing interests.

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Citation: Birhan M. 2019. Systematic review on avian immune systems. J. Life Sci. Biomed. 9(5): 144-150; www.jlsb.science-line.com

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