Brief Summary of Minutes of Annual Meeting

See attached document with NE-1016 minutes of October 2008 meeting in Washington, DC.

Objective 1: Identify and characterize genes and their relationships to disease resistance in poultry with an emphasis on the major histocompatibility complex as well as other genes encoding alloantigens, communication molecules and their receptors and other candidate systems.

A major focus of the multi-state project participants has been examination of genetic backgrounds of chickens with regard to susceptibility and resistance to various diseases plaguing the poultry industry. To that end, experimentation during this project has focused on major histocompatibility complex (MHC) haplotypes and selected genetic lines.

Studies at City of Hope (CoH) were devoted to defining the major histocompatibility complex (MHC-B and MHC-Y) in chicken in terms of nomenclature, gene content, genetic variability, and gene function. CoH completed a 242 kb map for MHC-B and obtained sequences for 14 haplotypes across a region of 59 kb from BG1 to BF2. CoH identified structural changes, synonymous and non-synonymous polymorphisms, insertions, deletions, and allelic gene rearrangements or exchanges that contribute to MHC-B genetic diversity. There is evidence among the haplotypes for whole and partial-allelic gene conversion events and for homologous recombination, as well as nucleotide mutations. CoH found that two CD1 genes map to the MHC-B. CoH defined the peptide binding preferences for two BF2 alleles showing that the BF2*21 allele found in the MHC-B commonly associated with Marek's disease resistance has a broad specificity, while the BF2*13 allele (often found in Mareks disease susceptible populations) has a far more stringent peptide binding preference. CoH also determined that MHC-Y class I molecules are widely and dynamically expressed in the spleen during development suggesting that these molecules may contribute to immunity in chickens.

Expression of B-MHC class I (BF1 and BF2) genes in several broiler haplotypes was evaluated by RT-quantitative PCR analysis at Auburn University (AL). BF2 was expressed in greater amounts than BF1 gene in all haplotypes, with a BF2 to BF1 ratio ranging from about 2:1 to 5:1. One exception was a haplotype in which BF1 transcripts were present in very low amounts, due to a mutation in the 3' splice site of intron 7.

AL evaluated polymorphisms in two genes of the innate immune response (Mx and OAS) in 14 commercial broiler breeder chicken lines from two different companies. A single nucleotide polymorphism (SNP) in the chicken Mx gene was reported to be associated with differential resistance to cellular infection with avian influenza virus (AIV) in cells ectopically expressing different Mx alleles. AL found that commercial broiler breeder lines had very low frequency of the Mx SNP associated with AIV antiviral activity, and some lines were fixed for the Mx allele that lacks antiviral activity.

In collaboration with Iowa State University (IA) and one of the commercial broiler breeder companies, AL evaluated associations between BF2 alleles in sires, and phenotypic traits of economic interest in their progeny. Associations were found between BF2 allele and some traits, including vaccinal antibody titers against infectious bursal disease virus (IBDV) and body weight. In the same study, associations between Mx SNPs in the sires and progeny traits indicated significant associations between Mx SNP determining differential antiviral resistance in the sires, and progeny traits including leg defects and early mortality.

In collaboration with the Southeast Poultry and Research Laboratory (Kapczynski and Suarez) AL evaluated the association of the Mx SNP encoding residue 631 in the Mx protein with differential antiviral activity on resistance to avian influenza virus in vivo. Results in two different commercial lines demonstrated that Mx Asn631 variants were associated with enhanced resistance (measured by mortality, morbidity, viral shedding) to a highly pathogenic AIV strain compared with Mx Ser631 variants. The Mx 631 variants were also associated with different cytokine responses in birds infected with AIV.

Specific-pathogen-free (SPF) flocks of chickens with defined MHC antigens are essential tools for the study of pathogens in the context of specific genetic backgrounds. Cornell University (NY) has maintained two MHC-defined SPF lines: the P2a (MHC: B19B19) and N2a (MHC: B19B19) which are susceptible and resistant to Marek's disease (MD), respectively. A major problem with the maintenance of commercial and research flocks of SPF chickens is infection with chicken infectious anemia virus (CIAV). NY has learned that CIAV can establish latent infections in the gonadal tissues of SPF chickens in the presence or absence of virus neutralizing antibodies and that CIAV can be transferred most likely as viral DNA from the hen to the offspring through the embryo. Viral DNA can be detected in many but not all organs of individual embryos suggesting that the virus is not transmitted through the germ line. Transmission can occur in embryos derived from antibody positive as well as from hens that are antibody negative hens. This type of transmission can also occur in commercial broiler lines. Studies on the transcriptional control of CIAV revealed that estrogen upregulates transcription but that COUP-TF1 and delta-EF1 down-regulate transcription. Thus viral replication occurs when the balance between positive and negative transcriptional controls is pushed toward up-regulation by estrogen and perhaps other hormones. These findings are important because these explain the difficulty faced by the SPF industry which often finds flocks seroconverting when birds are in full production.

At the University of New Hampshire (NH), six congenic lines containing B complex recombinants (R1-R6) on the Line UCD 003 background were tested for their primary and secondary antibody responses to SRBC. Lines with R5R5 (BF21-G19) and R6R6 (BF21-G23) had higher primary total antibody titers. Both secondary total and ME-resistant antibody titers were higher in R5R5 chickens compared with the other lines. The two recombinants having haplotype B21 had higher antibody responses to SRBC.

Six congenic lines containing B complex recombinants (R1-R6) on the Line UCD 003 background were tested for their responses to Rous sarcomas. The R1R1 (BF24-G23) and R4R4 (BF2-G23) genotypes had significantly higher TPI than the R2R2, R3R3, R5R5, and R6R6 chickens. Differences among B-F2 recombinants R2R2, and R3R3 versus R4R4 indicate that these three serologically similar recombinants possess different genes affecting tumor outcome. The comparable TPI between R5R5 (BF21-G19) and R6R6 (BF21-G23) suggests no effect of the B-G region.

Among chickens segregating for B1 and B2 haplotypes, B1B2 and B2B2 had lower tumor growth and TPI compared with B1B1 birds. Matings of B1B5 parents revealed lower tumor growth and TPI in the B1B5 genotype than was found in B1B1 or B5B5 homozygous progeny. This result indicated complementation between the more progressive B1 and B5 haplotypes. Subgroup C Rous sarcoma virus tumors grew differentially in progeny segregating for haplotype B1 in combination with either B2 or B5. B1B1 chickens had a TPI higher than either B1B2 or B2B2 chickens. The B5B5 chickens had significantly higher TPI than B1B5 or B1B1 chickens. B1 had a poorer response than B2 but was a better responder than B5.

The LEI0258 microsatellite marker had consistent allele size across multiple sources of the same MHC haplotype. In addition, some serologically distinct MHC haplotypes shared a common LEI0258 allele. Allele size varied due to internal repeats, plus a deletion.

Genotype R13R13 had lower primary total and ME-resistant antibody titers to SRBC compared with genotypes R13B17 and B17B17. Secondary total and ME-resistant antibody titers against the same antigen were again lower in R13R13 chickens compared with R13B17 chickens. The intermediate secondary titers for B17B17 chickens did not differ from either R13B17 or R13R13 chickens. Differential subgroup C Rous sarcoma virus tumor growth was found in congenic recombinant lines (R1, R2, R4, R5, R13). Genotype R5R5 with B-F/B-L21 had lower mean TPI than the R2R2, R4R4, and R13R13 chickens.

Congenic lines bearing serologically similar major histocompatibility (B) complex recombinant types R2R2 and R4R4 differed greatly in their susceptibility to MD. The incidence of MD was 19% in the 003.R2 line and 47% in the 003.R4 line (P<0.0001). The crossover breakpoints are separated by less than 1934 bp and they identify a single gene, BG1, as the locus affecting the observed difference in MD incidence in these lines. The two BG1 alleles are identical in coding region, but differ in 3'-untranslated region (3'UTR). BG1 encodes a receptor-like molecule containing an immunoreceptor tyrosine-based inhibition motif (ITIM).

In a broiler population at IA, microarray and quantitative RT-PCR assays were used to identify genes that have different transcriptional profiles associated with resistance to IBDV infection, as measured by viral load in the bursa. Seven genes were found to be co-upregulated only in resistant, but not in susceptible or mock-challenged, birds.

Global transcriptional analysis was used to investigate the genes associated with host response to Salmonella infection. Results on multiple genetic lines suggest that different chicken lines utilize different defensive mechanisms against the same bacteria. Results on cecum and spleen samples from an advanced intercross line F8 generation revealed many genes in immune response and signal transduction pathways to be differentially regulated after Salmonella infection.

Use of a high-density (3K) SNP panel and F8 birds was used to fine-map regions associated with Salmonella colonization levels. In total, 21 significant QTL regions were identified, in or near 19 novel candidate genes. Additionally, SNP in 13 of the beta defensins genes were genotyped in the same population, and five of the 13 genes were associated with bacterial burden. Two cross-over points in the beta-defensin cluster of genes were identified. Identifying genes involved in the host response to viral infection is not normally considered part of the immune response. As part of this effort at North Carolina State University (NC) has described changes in the expression and localization of ion transporter proteins in the intestinal epithelium following infection of turkey poults with turkey astrovirus.

Objective 2: Identify, characterize and modulate environmental and physiologic factors that regulate or affect immune system development, optimal immune function and disease resistance in poultry genetic stocks.

Work done at the University of California-Davis (UCD) determined the priority of leukocyte populations for key anabolic nutrients by examining the types and amounts of transporters expressed. Bursal cells have a high priority for glucose, branched-chain amino acids and lysine, but thymic cells have a very low priority relative to other types of leukocytes and most other tissue types. Furthermore, in the face of a deficit, bursal cells upregulate their ability to obtain glucose and amino acids, whereas thymocytes downregulate their uptake. Thus, the thymus is very sensitive to periods of food deprivation, energy deficiency, or amino acid deficiency, which cause a rapid decline in cellularity and weight. Fewer CD4+ T cells result in lower IgG production, while fewer CD8+ T cells result in diminished delayed-type hypersensitivity. For nutrients that the immune system is most vulnerable due to a low priority for acquisition, requirements based on maximal weight gain or egg production are likely to be inadequate for optimal disease resistance. Research was also initiated to identify key non-anabolic nutrients that serve as regulators of immune responses. It was found that nutrients that have strong immunomodulating activities include long chain polyunsaturated fatty acids, carotenoids, secondary plant compounds (e.g. genistein, flavonoids and many others found in herbals) and vitamins A, C, D, and E. These nutrients modulate the immune system by serving as ligands for nuclear receptors, including RXR and PPAR. The dose response relationship of these nutrients is non-linear and there are many unpredictable interactions between nutrients on immunity.

At NC efforts in two areas have been carried out in support of this objective. The first effort was focused on characterizing the innate immune response to viral infections and identifying gene expression profiles associated with enhanced resistance to disease. As part of this project NC sequenced and characterized the full length genomes of 8 turkey astrovirus isolates from commercial turkeys. This represents the largest catalog of astrovirus genome sequences of any species and is a critical first step in the identification of putative vaccine candidates. The second effort has examined the effect probiotic treatments on the immune response of broilers. NC has reported probiotic administration may inhibit the expression of pro-inflammatory cytokines and increase anti-inflammatory cytokines in the intestines of developing broilers. In addition, probiotics administration also appears to modulate the amount of energy consumed by the immune system which may help explain field observations that probiotics promote animal health.

Both CIAV and MDV can modulate immune functions. CIAV can reduce or eliminate specific cytotoxic T-lymphocyte (CTL) responses to other pathogens. NY has shown that concurrent infection of chickens with CIAV and REV can reduce or eliminate CTL responses to REV at 7 days post-infection. The focus of Marek's disease studies was the importance of specific genes for the pathogenesis of MD. Using specific deletion mutants of the very virulent RB-1B strain we found that deletion of R-LORF4 resulted in attenuation of MDV with a very low tumor incidence at 13 weeks of age in contrast to the wild-type RB-1B Bac-derived virus which caused tumor in 100% of the chickens within 6 weeks post-infection. Likewise, deletion of exon 1 of vIL-8 reduced the pathogenicity significantly. Although the mechanistic reasons for these effects are not fully understood the findings will be important for the complete characterization of the virulence factors of MDV and may lead to the development of safe and more effective vaccines. NY also found that the very virulent plus (vv+) strains caused a severe proinflammatory response in the spleen and the brain of susceptible P2a chickens and especially in the resistant N2a chickens causing increased mortality in the absence of tumors. These results indicate that breeding for increased resistance to MD tumors may not necessarily increase the resistance to other pathologies associated with MDV infection.

The ultimate goal of research at Western University (WU) is to exploit mechanisms that enhance viral specific CD8+ T lymphocyte immunity to improve the efficacy of recombinant vaccines. Studies on low path N5 AIV infection in B2B2 and B19B19 chicken lines (Northern Illinois University) found that: 1) memory CD8+ T-cell populations are induced in vitro with specific APC in an MHC restricted manner; 2) memory CD8+ T-lymphocytes in chickens express higher levels of CD44 than naïve T-cells; and 3) fluorescence intensity of CD44 expression is greater on memory CD8+ than CD4+ T-cells.

The DNA vector study demonstrated that both HA and NP of AIV stimulate CD8+ T-cell responses. However, the response with NP responses was significantly greater than that with HA. The non-replicating adenovirus vector induced HA responses. An effector CD8+ T-cell response was observed at 10 days post-infection with the HA vector and the CD8+ memory responses was observed by 3-5 weeks post-infection. Thereafter, similar to the DNA plasmid and low path AIV, the response declined. A booster was given in this study, which induced a rebound of the T-cell response. The rebound was equal to but not significantly greater than the initial primary response and also decreased with time.

A previous study on REV infection of prairie chickens conducted at WU showed that a DNA vaccine expressing the REV envelope protein (env) did not protect infected chickens from disease. Based on reports that the tegument protein VP22 of herpesviruses fused to the antigen can increase the immunogenicity of DNA vaccines, three groups of Attwater's greater prairie chicken hybrids were inoculated with: 1) DNA plasmid containing 100 mg VP22; 2) DNA plasmid containing 50 mg VP22/ENV and 50 mg VP22/GAG; and 3) PBS. Vaccination of prairie chickens with a DNA vaccine did not prevent infection with REV, despite use of large doses of DNA and a comparatively low infectious dose. Analysis of PBMC collected from both uninfected and chronically infected Attwater's prairie chickens for REV protein expression (gag) in lymphocyte subsets revealed that between 50-64% of the total lymphocytes were infected with REV. Though vaccination did not prevent infection, vaccinated birds showed lower percentages of infected lymphocytes, including CD4+ cells. One third of the infected lymphocytes did not express CD4 or CD8. Percentages of CD4 cells decreased in naturally infected birds, whereas those of CD8+ cells increased in both naturally and experimentally infected birds, indicating some form of control of infection.

Using the Smyth line, which spontaneously develops autoimmune/autoinflammatory disease (specifically vitiligo) and control lines of chickens, the cause-effect relationship between a genetically controlled disease, immune function, environmental factors and gender were investigated at Arkansas (AR). Approaches used included: gene expression analyses, especially cytokine and chemokine expression in the autoimmune lesion prior to and throughout the development of vitiligo; in vivo and in vitro analysis of antioxidant capacity and oxidative stress; examination of the role of herpesvirus of turkey and other environmental triggers in the expression of vitiligo in genetically susceptible individuals. Through these studies it was established that: there is an inherent melanocyte defect in Smyth line chickens including altered antioxidant capacity and lipid peroxidation, the anti-melanocyte response is a T helper cell type 1 polarized cell-mediated response that is accompanied by melanocyte-specific autoantibodies, and that inflammation may serve as a trigger in the expression of Smyth line vitiligo. Genetic analyses looking for candidate genes involved in vitiligo susceptibility are underway in collaboration with researchers in Sweden.

Other contributions by AR include studies on pulmonary hypertension syndrome (ascites) in broilers revealed that genetic susceptibility to this disorder is reflected in the broiler's ability to generate innate immune activity, particularly the appropriate balance of vasodilatory and vasoconstrictive factors. To gain insight into cellular immune response activities, leukocyte infiltration in response to injection of primary and recall antigens into integumentary tissues was examined in chickens. Histological and immunohistochemical analyses revealed a leukocyte response pattern that was chronologically, qualitatively and quantitatively similar to that described in mammals. These studies have also led to the development of a new procedure to examine and monitor cellular or tissue immune response activities in birds using the growing feather as an in vivo test tube. Lastly, AR also examined the transfer of maternal antibodies from the hen to the egg and subsequently to the chick, providing a better understanding of this protective mechanism important in the first weeks of a chick's life.

Objective 3: Develop and evaluate methodologies and reagents to assess immune function and disease resistance to enhance production efficiency through genetic selection in poultry.

A category of crucial biological materials needed for the successful conduct of research on genetics of disease resistance is specialized genetic lines and resource populations of chickens. IA maintained many very highly inbred lines of birds, including several sets of MHC-congenic lines. These lines were used to assess the impact of host genetic variation on immune response and disease resistance, both at IA and elsewhere, with collaborators. Approximately 2000 chicks were hatched per year. Semen samples from all roosters of all inbred lines were collected for two generations and sent to the US Germplasm Preservation Lab in Ft. Collins, CO.

In addition to the pure lines, a mapping resource population was developed to aid the identification of genes related to immunity and disease. The Iowa Salmonella Response Resource Population (ISRRP) was produced from an initial cross of outbred broiler males with females from the highly inbred Leghorn and Fayoumi lines. Thereafter, the birds were intercrossed each generation to propagate advanced intercross lines (AIL). The F8 AIL generation was expanded in bird number and phenotyped for response to challenge with pathogenic Salmonella enteritidis (SE) or to antibody response to vaccination against SE. Tissue samples from this generation were then used for experiments, as reported under Objective 1.

Primer pairs and optimal conditions for assessing mRNA expression levels were determined for about 30 genes putatively related to immune function. They were used to determine gene expression in tissue samples from pure lines or AIL F8 birds that were challenged with Salmonella. The expression of several cytokine and other genes was found to be associated with host response to bacterial infection.

Collectively, the research conducted has revealed new candidates genes for host resistance to viral and bacterial diseases in poultry, which can be used as a foundation for additional studies or applied in marker-assisted selection to enhance innate resistance to disease.

Over the last few years, the University of Guelph (UG) has been able to develop a low-density immune system microarray and has applied that to investigate immune system gene expression in lymphoid tissues of chickens infected with MDV, especially chickens of genetically resistant or susceptible lines. Furthermore, UG have characterized cytokine responses to MDV infection in several tissues that play an important role in the pathogenesis of this virus, such as feather tips and bursa of Fabricius. UG has confirmed that the virus elicits host responses in feathers leading to cellular infiltration and cytokine production. However, these responses are not effective and the virus can still be shed from chickens that are vaccinated and protected against Mareks disease. The above findings have shed more light on understanding the process of MDV pathogenesis and immune response to this virus.

AL developed a molecular method to type BF2 (MHC class I) alleles, by PCR amplification with locus-specific primers followed by nucleotide sequence determination. Also, AL developed a rapid method to type Mx alleles for the SNP associated with differential antiviral activity, by PCR-RFLP. Recombinant core proteins of REV were generated at WU with a His plasmid and purified with a nickel column. The protein was used to make antibody specific for REV. This antibody was used to detect infection using flow cytometry and IFA. Detection of REV infection in lymphocytes of infected birds using IFA or flow cytometry detected infection where the nested PCR did not. Therefore, detection of cellular presence of REV protein was a more sensitive indicator of REV infection. A nitric oxide (NO) assay was developed to detect ex vivo activated and in vitro stimulated T cells. Flow cytometry was developed for evaluating effector and memory T-cells using MAb specific for chicken CD44 antigen with MAb specific for either chicken CD8 or CD4 antigens.

Development and characterization of reagents to be used to study the turkey immune system were done at NC. NC has tested a large panel of chicken specific monoclonal antibodies for cross-reactivity with commercial turkeys and identified 8 antibodies which demonstrate reliable cross-reactivity. NC has developed RT-PCR and Real-Time RT-PCR reagents for approximately 19 turkey genes. In addition NC has developed peptide antibodies specific to the turkey iNOS gene as well as developed a turkey specific bioassay for IL-8. AR developed a minimally invasive in vitro method to monitor immune/tissue responses in an individual by injecting antigens and other test material into the pulp of growing feathers.

The examination of inflammatory mediators in the chicken Harderian gland and circulating thrombocytes has been the focus of work conducted at the SC station. Hematopoietic-prostaglandin D2 synthase (H-PGDS) expression was a good indicator during the early stages of immune responses of reactivity to the Infectious bronchitis/Newcastle disease vaccine. H-PGDS is present in many different types of cells in the body, and PGD2 is becoming recognized more as an important chemoattractant/activator modifying activity of cells involved in both pro- and anti-inflammatory actions. Considering the relatively small literature base on chicken thrombocytes, additional knowledge gained on this cell's ability to be activated and produce bioactive molecules will improve our understanding of its role in immunity. These cells are nucleated and thus possess a full complement of genetic material capable of being induced for expression of certain immune factors. The increasing uniqueness of the thrombocyte is putting it into a different light of interpretation regarding active, overt participation in innate immunity. Data collected (SC) found thrombocytes to express many of the same innate attributes and functions of macrophages and heterophils. Chicken thrombocytes possess TLR4 that is connected to cytoplasmic signaling pathways leading to gene expression. Furthermore, the differential influences of MAPK and NF-kB pathways on expression of IL-6 and COX-2 (and IL-8 via BMS345541 inhibition) are apparent in inhibitor treated thrombocytes. Cytoplasmic signaling pathways in thrombocytes are devoted to immunologic responses in support of inflammation.

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