Annual/Termination Reports:

[02/27/2003] [12/08/2004] [11/19/2007] [06/06/2007] [03/04/2009]

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Brief Summary of Minutes

Accomplishments

OBJECTIVE 1. Several NE-60 stations (CA, IA, NC, NH, NYC, NIU, ARS-ADOL) maintained important genetic resource populations despite budgetary constraints. <br /> Attempts were made to identify MHC haplotypes in commercial broiler lines. Genotyping was done using restriction fragment length polmorphism (RFLP) for B-G, and RT-PCR followed by cloning and nucleotide sequence determination for B-L and B-F genes (AL). Eight haplotypes were characterized in Line C, and two haplotypes in Line E (AL). In combination with the 10 haplotypes previously characterized in broiler breeder Line A, seven haplotypes were identified that encoded novel B-F and B-Lb alleles not previously reported in Leghorns (AL). The influence of two MHC haplotypes (BA4 and BA5) on disease resistance in broiler breeder Line A was also tested (AL). BA5 haplotype was associated with relative resistance to cellulitis induction by E. coli, by comparison with BA4 haplotype. The reverse was seen in collaborative studies (AL and DE) in which chicks were challenged with RB-1B strain Mareks disease virus (MDV). DE has characterized the molecular MHC genotypes in a commercial broiler population. At least 4 class II and 8 class IV RFLP genotypes were discovered whose frequencies varied among three lines within the population. At least one more class II allele that was not previously reported was identified. An MHC gene, B11, that appeared to confer a significant degree of MDV resistance was studied by CA. <br /> Eight non-MHC alloantigen systems were examined for their effects on chicken mononuclear phagocytic system cells (NC in collaboration with NIU). Chicks of P allotype carrying P13 and P1 allele were high responders for LPS-induced nitric oxide synthase (iNOS) expression and activity, whereas macrophages from I8 allotype chicks produced highest levels of IL-6. This suggested that alloantigen systems can exert significant influence on immune endpoints independent of the B complex (NC). Non-MHC alloantigen systems had significant effects on Rous sarcoma virus (RSV) tumor outcome in a progeny of matings between B2B5 sires and B5B5 dams derived from a cross of Modified Wisconsin Line 3 and Line NIU 4 White Leghorns. (NH). <br /> The influence of B and L haplotypes on immunity to E. tenella was evaluated. The results support prior studies showing lower lesion scores in the B5B5 genotype compared to B2B2 chickens (NH).<br /> Identification of candidate genes for response to Salmonella enteritidis (SE) was of interest to IA. Associations of the following genes were found with one or more traits of SE response: MHC class I, NRAMP, IAP1, caspase, and prosaposin (IA). <br /> Several groups also examined gene systems with involvement in immunocompetence. A chicken genomic DNA clone was found to contain a region of chicken chromosome 1 and it was syntenic with human chromosome 22 (DE). This clone was determined to contain the genes, Synapsin 3 and FBX07. To identify other non-MHC genes that affect MD resistance in broilers, DE established an F2 population, challenged it with RB1B, and recorded MD clinical signs, lesions and mortality. <br /> A collaborative project between (DE) and ARS-PBESL sought to identify genes affecting coccidiosis resistance. At least one quantitative trait locus (QTL) was identified on chromosome 1. Furthermore, a genetic component was identified (DE) that influences body weight during a MDV challenge in broilers<br /> Other studies compared the iNOS expression and activity between B15B15 (Cornell K-strain), B6B6 (G-B2) and B13B13 (G-B1) genotypes (NC). Macrophages from these chicken lines can be hyper- or hypo-responders for iNOS gene expression and activity regardless of the source of bacterial lipopolysaccharide. <br /> NH studied major histocompatibility (B) complex effects. The outcome of Rous sarcomas was assessed in second backcross generation BQB17 chickens with a genomic composition of 87.5% inbred Line UCD 003 and 12.5% inbred Line UCD 001. The results of this study suggested complementation between the BQ and B17 haplotypes.<br /> <br /> OBJECTIVE 2. PEMS is a multifactorial disease of turkeys that is characterized in part by immune dysfunction. NYC and NC have isolated of a novel reovirus (ARV-CU98). This astrovirus induced significant growth suppression, diarrhea, thymic and bursal atrophy and altered B-, T-, and macrophage functions. <br /> The finding that chicken infectious anemia virus (CIAV) was present in SPF chicken flocks had a major impact on NYC research. In order to generate quantitative data on the effects of experimental infection with CIAV on CTL generation, NYC developed real-time, Taqman-based, PCR and RT-PCR assays. These assays can also differentiate between two CIAV strains, Cux-1 and CIA-1. NYC found that CIAV can be an important immunosuppressive agent that can modulate immune responses.<br /> Polychlorinated biphenyls (PCB) are important environmental contaminants. Chickens treated in ovo with 0.39 ng/g 2,2-dichlorobiphenyl (2,2¢-DCB) had significantly lower primary antibody titers against SRBC compared with the vehicle-treated and control chickens (NH).<br /> Dietary factors were shown to have significant impact on performance and immunocompetence of chickens. Mycotoxins were shown to significantly suppress several immune parameters in direct-fed chickens or in progeny chicks (NC). Dietary addition of other compounds such as beta-glucans improved various immune endpoints in chickens (NC).<br /> AR also conducted several studies on nutritional supplements demonstrating beneficial immunomodulatory effects of dietary supplementation with vitamin E or bacterial cell wall products from both Gram + and Gram- bacteria.<br /> FRAMINGHAM revealed that including mannanoligosaccharide in poultry diets increases levels of both natural and acquired antibody. <br /> The importance of nitric oxide (NO) as a modulating factor for Mareks disease was examined. The most likely cells producing NO are macrophages or their precursors present in the CEF cultures. In most experiments, N2a chickens showed an increase in plasma NO levels at 7 dpi compared to the susceptible lines P2a. The very virulent (vv)+ strains of MDV caused significant higher levels of NO in the plasma than the less virulent strains. NYC also examined the importance of rIFN-a for the activation of natural killer (NK) cells by inoculating chickens with rMDV- IFN-a or by feeding rIFN-a in the drinking water. In both instances IFN-a caused a decrease in NK cell activity in spleen cells.<br /> AR focused studies on the mutant Smyth line chicken which exhibits genetically controlled spontaneous autoimmune vitiligo. AR has demonstrated that pigment cell death in Smyth line chickens involves an inherent melanocyte defect, an environmental factor , and immunological factors. <br /> One study evaluated primary and secondary antibody response to BA and SRBC in eight different genetic groups consisting of four different MHC types on two different line backgrounds. The overall results indicated complex interactions between specific MHC alleles and the non-MHC background of the lines in which they were studied (IA).<br /> Other experiments aimed at identifying genomic regions and specific genes controlling antibody kinetics in adult hens. Several candidate gene fragments were sequenced, including MHC class I and II, IgL, IAPI, ZOV3, IFN-g, TGF-b2 and b3 (IA).<br /> Thyroid hormone and thymulin were studied extensively by NYC. The station found that low level (0.1 ppm) thyroid hormone (i.e., triiodothyronine or T3) supplementation enhances both IL-2 activity from culture supernatants and IL-2R expression while higher levels (1.0 ppm) decrease the expression of both of these activities. Thymulin was found to modulate both the percentage of CD4+ and CD8+ cells and the mean fluorescence intensity (MFI) of these co-receptors. (NYC). The initial thyroidal status of a strain or of an individual appears to correlate with baseline natural killer (NK) cell activity. T3 supplementation may enhance NK cell activity, particularly in the T3-deficient SLD and this appears to be at least partially due to an enhanced responsiveness of NK cells to IFNg (NYC). Dietary T3 supplementation significantly increased abdominal exudate cell (AEC) responsiveness to rcIFNg in the hypothyroid SLD strain while not affecting AEC responsiveness in the K strain. T3 levels affect avian IFNg activity by influencing both the responsiveness to and the production of IFNg. Direct in vitro exposure of chicken splenocytes to thymulin suggests that one function of thymulin is to enhance the responsiveness of these cells to IFNg, perhaps by enhancing the expression of IFNg receptors. The ability of thymulin to act directly on macrophages was examined by testing the effect of in vitro thymulin treatment on HD-11 (a macrophage cell line) responsiveness to rcIFNg (NYC). rcIFNg treatments alone consistently increased NO production in a dose-dependent manner. <br /> NYC most recent work has focused on the ability of thymulin to stimulate in vivo NK responsiveness to a viral infection and suggests that we can enhance the resistance to infection with avian infectious bronchitis virus (IBV) with short-term thymulin supplementation.<br /> Matrix metalloproteinases (MMP) were examined in RSV tumors from NH congenic lines 6.6-2 (B2B2) and 6.15-5 (B5B5). B5B5 birds had significantly higher mean tumor profile index (TPI) than did B2B2 chickens. Congenic Line 6.15-5 tumors had significantly more RSV tumor sections that stained positively for MMP-2 and MMP-9 than did congenic line 6.6-2 tumors. <br /> PA investigated the role of cytokines and hormones involved in the initiation of humoral immunity, using lipopolysaccharide (LPS) [T-independent antigen] and bovine serum albumen (BSA) [T-dependent antigen]. It was concluded that although LPS and BSA injection can induce a humoral antibody response in chickens, the mechanism of antibody response initiation involving cytokine network and neuroendocrine system activation are different for each antigen. PA also found a possible recirculation of lymphocytes from circulation to different lymphoid compartments during septic response and to the local site of injection during the cutaneous hypersensitivity response. In studying the role of the photoperiod and melatonin on broiler performance and on the immune responses, PA demonstrated that using intermittent lighting (increasing the dark period), is advantageous over using 23 hrs of light for enhancing both production performance and immune responses. <br /> <br /> OBJECTIVE 3. Non-gene-specific genetic factors related to the response to Salmonella enteritidis (SE) in young chicks were investigated. In one experiment, using layer chicks, line was demonstrated to be highly significant on survival. In surviving chicks, the genetic line effect was significant on cecal SE burden but not on spleen, and these two measures were under independent genetic control (IA). In broiler breeder chicks, a significant negative correlation of vaccine antibody response and bacterial burden in cecum indicated that chicks with greater genetic potential to respond to vaccine are better able to control and eliminate the pathogenic bacteria (IA). <br /> Genomic regions associated with vaccine or challenge response to SE were identified in a resource population formed by crossing broiler sires and inbred (Leghorn or Fayoumi) dams. Microsatellites were used as the marker system. Associations of four microsatellites were confirmed for vaccine response and SE bacterial burden in spleen and cecal content (IA). <br /> The AL station developed a PCR with sequence-specific primers (PCR-SSP) to type alleles of the B-LbII locus. Serological reagents to identify new haplotypes discovered in broiler breeder chicken lines were produced. Another project was development of quantitative measures of bursal damage and viral load in chickens challenged with infectious bursal disease virus, for the purpose of assessing MHC associations with disease resistance. Finally, analysis demonstrated that broiler MHC haplotypes varied dramatically in expression of MHC class I molecules on lymphocyte cell surface; furthermore, MHC class I expression correlated with proliferative response to Concanavalin A. <br /> To map quantitative trait loci (QTL) associated with disease resistance to avian coccidiosis and growth, two commercial broiler lines with different degrees of resistance to the disease were crossed to generate F1 generation that were intercrossed to produce 314 F2 generation offspring (ARS-PBESL). Inter-trait comparisons revealed that oocyst shedding was a good parameter for evaluating disease resistance/susceptibility. A locus on chromosome 1 associated with oocyst shedding (LOD = 3.46) was revealed. The genetic mechanism of this locus appeared additive. The genomic scan also identified three potential growth QTL on Chromosomes 1, 6, and 8 (ARS-PBESL). <br /> NYC developed an approach to study CMI responses to MDV using the REV-based system. MHC-defined REV cell lines that can be lysed by syngeneic REV-sensitized CTL were stably transfected with MDV genes and used as target cells for the detection of MDV-specific CTL recognizing gB, pp38, meq, and ICP4 epitopes, although the latter one was only detected using CTL from the resistant N2a line. The effector cells were characterized as CD8+CD4- CTL expressing TCRab1. During the current NE-60 project period, NYC found that REV cell lines expressing the immediate early protein ICP27 are lysed by syngeneic effector cells from both P2a and N2a chickens. More recently a large study was completed in which CTL responses to MDV glycoproteins gC, D, E, H, I, K, L, and M were examined. Cell lines expressing gI were lysed by effector cells from N2a and P2a chickens, while N2a-derived CTL also lysed cell lines expressing gC and gK and to a lesser extent gH, gL, and gM. P2a-derived effector cells did not lyse cell lines expressing these glycoproteins, but did lyse cell lines expressing gE. The importance of these studies is that the differential recognition of MDV proteins by CTL from chickens with different MHC genes contributes to the understanding of differences in genetic resistance to MDV. <br />

Publications

Impact Statements

1. Understanding of the structure, function and disease-resistance relationships of both MHC and other genes has seen substantial progress during the current project period. Examination of inbred, non-inbred and congenic populations further defined MHC and non-MHC effects on specific diseases including MD, Rous sarcomas, Salmonella enteritidis, Eimeria tenella, E. acevrvulina, and CIAV. In addition, genetic influences on basic immunological functions were defined. Use of unique MHC region recombina
2. Characterizing environmental and physiologic factors that affect the immune system progressed in the current project period. Immunosuppressive effects of PEMS and CIAV pathogens were studied. Exogenous administration of thyroid hormone, thymulin, and IFN-g had immunomodulatory effects. Thymic-endocrine interaction and endocrine function of the thymus were studied. An inherent melanocyte defect, an environmental factor, and immunological factors were identified as important in chickens exhibiting
3. Genomic regions as well as non-gene-specific genetic factors related to the response to Salmonella enteritidis were described for layer and broiler breeder chicks. Whole genome scans using microsatellite markers and variance-component linkage analysis identified QTL associated with production and disease resistance. This analysis found one QTL for resistance to coccidiosis and three QTL affecting growth. A PCR test was developed to type B-LbII alleles. Lymphocyte MHC class I expression was found

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Brief Summary of Minutes

Accomplishments

Extensive collaboration among members was evident and much meeting discussion focused on genetic resources and conservation of living stocks. A special tribute to Dr. W. Elwood and Mrs. Ruth Briles was held in conjunction with the meeting and included an evening dinner celebration. The tribute recognized the enormous contributions of the Briles to the NE-1016 (formerly NE-60) group and to the field of avian immunogenetics. <br /> Studies continued to elucidate the molecular and cellular basis for resistance and susceptibility to Mareks disease, Salmonella, Rous sarcoma virus, autoimmune disease, and general immune function. Much progress was made in profiling of genes involved in resistance, exploration of new genes and determination of the contribution of genetic variation to disease resistance and susceptibility. Work continues in these areas. <br /> <br /> Auburn University (AL). Quantitative differences in mRNA expression of BF1 and BF2 loci alleles for broiler breeder lines were established. BF1 of four haplotypes have the same sequence of alpha 1 and alpha 2 regions, one haplotype has an additional nucleotide difference leading to two different amino acids. Locus specific probes were designed for BF1 and BF2 for real time qPCR. Transcript differences were evident and closely related BV1 alleles may exhibit wide differences in expression; regulatory elements could contribute. A new area of research is the analysis of the functional polymorphism in the Mx gene. Mx has antiviral function (mice) and Mx1 possesses a major impact on resistance and susceptibility to influenza; Mx proteins block viral replication. A PCR test was developed to identify polymorphisms in the Mx gene.<br /> <br /> <br /> Beckman Research Institute (CA). Work continues on defining functions of B and Y regions of the MHC. Major sequencing effort with international collaborators continues. To date, 15 Ylec loci are found intertwined w/16 non classical Class I loci in 255 Kb. An overall objective is to search for the gene(s) impacting MD and tumor growth. Studies of Ylec molecules are in progress; these may be involved with NK cell activities and may be used by viruses as decoys to evade NK cell activation. Work continued on isolated peptides to establish features of the Class I binding site domains; amino acid motifs (dominant to infrequent) and models with folded protein structure were described. Motifs can be used within a web-based alogorithm to search what other proteins (and their sources) have similar motifs, e.g., from RSV and MDV (candidate proteins among viral proteins). Prior research showed that BR2 and BR4 recombinants, which are serologically identical, differ in Mareks disease incidence (BR2 incidence is 14% and BR4 is 48%) this was further confirmed and these were found to differ in crossover breakpoint defining a candidate gene region for influence of MHC in MD (UCD, Wakenell; UNH, Taylor).<br /> <br /> Cornell University (NY). SPF flocks were remarked on, and the priority over the next 5 yr is to conserve these including P2a (B19, C2, MDV susceptible), N2a (B21,C2 MDV resistant). Cornell also maintains a White Peking duck flock including 20 females 8 drakes. CIAV latency and reactivation studies continued. Four hormone response elements sites were identified in the promoter region along with a SP1 local site. These were tested for hormonal control, using various constructs, and transfections of DF-1, LMH, LMH/2A, theca and granulose primary cells. Downstream elements down regulate transcription, and long promoters resulted in fewer transcripts by qPCR. vvMDV impact on cytokine levels in the N2a and P2a SPF flocks was studied. Lines exposed to virulent JM-16 or vv+RK1 were examined and virus replication studied by qPCR, cytokine transcripts were measured (spleen and cerebellum), nitrous oxide (NO), and also histopathology for lesions. High responses were seen in the N (resistant line). Conclusion is that vv+ causes a strong proinflammatory response in brain, independent of MHC type.<br /> <br /> <br /> Clemson University (SC). Hardarian gland (HG) studies continued to establish cytokine expression with specific emphasis on IL-6 involvement using a bioassay for IL-6 activity. Proteomic analysis (S. Burgess) of HG supernatant fraction was conducted and protein sequences compared. Several proteins were cloned and sequenced including H-Prostaglandin GD2Synthetase. The C1 Vick line was studied and the enzyme system was found to be present. Studies contribute to understanding mast cell biology; intermediate and end products of mast cell metabolism (hypersensitive immune response) involves immediate and latent effects due to mobilization of arachidonic acid pathway and production of prostaglandins, PGE2 is of prime importance given most symptoms.<br /> <br /> <br /> Iowa State University (IA). Gene expression tools were established (microarray and qPCR) to study variation in the founder lines for the ISU resource population. Following Salmonella exposure, mRNA was examined at 2 and 18 hr, using chicken 13K cDNA array from the Hutchinson Cancer Research Ctr; spleen and cecum were studied. A mixed model approach (rather than threshold change) was the approach for data analysis. Applied consistency criteria to reduce number of genes to consider for qPCR as 100s of differentially expressed genes were identified at threshold at both times or for all lines. Genes included those one would expect, MHC class II up-regulated, Ig light chain down-regulated, in addition to novel genes such as quiescence specific protein precursor (A30230). The goal is to use data from advanced intercross (F8) to identify important genes, segregating SNPs, develop immunohistochemistry and proteomics profiling. ISU Genetic Resources at ISU for Chicken Biotechnology Research include the following: 20 inbred line, many MHC congenic sets; 4 partially inbred Leghorn line (B X IrGAT); 1 non-inbred broiler breeder male line; 2 adv intercross line (broiler x inbred) (now F8). Genetic line priorities include the Advanced Intercross, G-B1 and G-B2 Leghorn lines, M5.1 and M15.2, Fayoumi, a broiler line, and the lines from 1925 inbreeding program.<br /> <br /> <br /> Northern Illinois University (IL). An exacting search for recombinants involving crossover between B19 and B21 was conducted; over 1000 chicks were examined, but no recombinants were recovered indicating B system recombination is a rare event. Haplotyping of five Hy-Line commercial stocks was undertaken (closed WL female breeder lines) for A,B, C, D, H, I, L, P and M systems. On average 1-2 alleles were found per system (and three for P in one line). LEI0258 microsatellite marker was used for MHC allele detection (Fulton); B13 and B21 are serologically different but exhibit the same LEI0258 amplicon size; by SSCP a 1-2 nt difference was detected. Ring Necked Pheasants were dropped from the NIU collection, the Northern and masked bobwhite colonies remain. BL polymorphisms are being examined by SSCP to categorize patterns. The objective is to maintain 12 MHC haplotypes from the original colony (1999). Evidence was found for a trisomic condition in Line 4 for GGA 16. Typing is ongoing w/ non cross reactive alloantisera, and LEI0258 wherein three amplicon sizes were identified (195, 264 311 bp sizes for B1, B2/B29, B10). <br /> <br /> <br /> Texas A&M University (TX). Enhancement of immune response of commercial poultry to eimera species was studied. Mechanisms were explored for new vaccine strategies and sensitization. Three experimental strategies were explored: Drinking water antigen (BSA) exposure, CpG oligonucleotides, and enhancement via live oocyst vaccination. Increased IgG levels were dependent on route of administration. For protein antigens drinking water may be an effective route. Also examined immunity boost with non-methylated CpG oligonucleotides as an adjuvant to BSA using a single oral administration to explore the resulting Ig status. Results indicated that, given orally, CpG can increase Ig secretions. <br /> <br /> <br /> University of Arkansas (AR. Smyth line chicken is an animal model for human autoimmune vitilligo, a pigment disorder, wherein pigment is lost in patchesthis is the only animal model for the human disease. Recent work focused on the issue of whether there exists a defect in anti-oxidative capacity in the melanocytes and their ability to respond to stress. Smyth cultured cells were found to possess an inherent susceptiblity to lipid peroxidation (higher in Smyth cells) vs Brown Line (parental stock) or Light Brown Leghorn (control) lines. Catalase activity was similarly heightened in Smyth line cells. In vivo, in the feather tissue, heightened levels of lipid peroxidation correlated to onset of disease. Other autoimmune diseases are found in Smyth line, alopecia and autoimmune thyroiditiis (as in human).<br /> <br /> <br /> University of CaliforniaDavis (CA). A new survey is in progress for listing poultry and avian genetic research resources, to update that published within the Avian Genetic Resources Task Force Report in 1999. The information will be presented within a searchable website. Results were reviewed from a collaborative study (Miller, Bacon) exploring MHC expression in the early chicken embryo at 0-48 hr of incubation. Results indicate that B-F and B-L are expressed on cells of early differentiating embryos at each of the stages examined whereas B-G was found at only at one stage (12 hr blastula stage). These results show that the avian embryo, like the mammalian has MHC expression prior to the elaboration of the immune system.<br /> <br /> <br /> University of California-Davis (CA). Reviewed studies of MDV resistance of R2 and R4 birds after challenge w/ RB1B (vv strain of MDV). Serum chemistry (dried format) can provide good insight to health status of birds and provide correlates with typical MD phenotype (e.g., lesions, etc.). Liver enzyme analysis was utilized (alkaline phosphatase (AP)). Lesion profiles of R2 and R4 birds were typical (kidney, liver, gonad etc.) although thymus and lung tumors were observed (which are not typical). Interestingly, in the inbred/congenic lines bearing liver tumors, retrospectively a drop in AP levels could be detected three to four weeks prior and so one could pick up initial disease before the animal succumbs to virus. Such predictability is of great value for future. Note this drop in AP level is opposite to what happens in mammals with liver disease; the decrease remains to be explored.<br /> <br /> <br /> University of Delaware (DE). F-box proteins are involved in the ubiquitization of proteins which marks them for degradation (e.g., viral proteins). Database ESTs were searched to identify proteins for motifs and 20 were identified allowing the categorization of three classes. Analysis keyed on specific proteins that might be involved in cancer, tumor transformation, and viral presentation and a subset will be studied for their expression in immune-activated tissues. Proteins were characterized of MD-stunted birds (commercials) in families with high incidence of growth defect during MD challenge. Unusual proteins were identified in spleen including annexin 5 (involved in apoptosis, protein metabolism). DNA of birds was archived to retroactively examine for correlated alleles.<br /> <br /> <br /> University of New Hampshire (NH). Characterization continued of responses to RSV of the B1 haplotype from a commercial line. B1/B2 and B1/B5 haplotypes were tested by exposure to RSV subgroup D and tumor growth was monitored. Pattern of tumor growth and analysis of genotypes are underway. LEI0258, a microsatellite marker is being used as a marker to discriminate the haplotypes. Studies continued in producing congenic MHC recombinants on UCD 003 including R1-R6, and R13, R7, 10, 11, 12. Although for some lines, only created one cross w/UCD 003 and then discontinued due to financial constraints. <br /> <br /> <br /> Wagenigen Agricultural University (NL). Immunomodulation studies were conducted involving primary and secondary immunization w/HuSA. The Ig, IgM, IgG responses of various treatments of LPS, PBS, KLH were studied and it was observed that these homotopes enhanced responses, likely a non-antigen specific recall response. Non-steady state situations (complex stress simulations: starvation, long term cold stress) were examined to explore if chickens respond/adapt to these stressors. Antibody responses were not affected, specific antibody responses were normal; however, natural antibody levels were affected by cold stress. Over time, adapted responses were observed for continuous cold stress (using both high and low antibody lines). Body weights were reduced, at half rate of expectation. Studies show that immune response has a high priority as compared to growth. Research speaks to homeostasis: IR has low energy requirements (as compared to growth), but gets priority under stressed situations. <br /> <br /> <br /> Wagenigen Agricultural University (NL). Studies of QTLs for KLH response in commercial feather pecking population (behavioral differences) indicated three QTLs detected on GGA 12, 14, 16. GGA14 (~80 cM) fine mapping was conducted to find candidate genes using SNP markers in one family. The QTL was narrowed toward the end of the chromosome, with the greatest effect at the end of the available markers, ~78/80 cM. In early generations a paternal allele effect was observed i.e., parent of origin effect on the QTL, in downstream generations observed lack of effect when allele was inherited from female in downstream generations. Results provide preliminary evidence for parent of origin effect for immune traits. <br />

Publications

Impact Statements

1. Substantial new detail is available from research outline expression of genes involved in immune function in tissues of chickens showing resistance and susceptibility to disease.
2. Sequencing and cloning of novel genes and QTL regions continues with support from the newly available draft chicken genome sequence.
3. Genetic resources are being maintained in the face of decreasing budgetary support at the institutional level and this remains a critical area needing a solution.
4. Interactions between research groups remains at a very high and productive level.

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Participants

Members present: Erf, Gisela (gferf@mail.uark.edu) - University of Arkansas; Ewald, Sandra (ewaldsj@vetmed.auburn.edu) - Auburn University; Heidari, Mohammed (heidairi@msu.edu) - USDA, ADOL; Kopulos, Renee (ebriles@niu.edu) - Northern Illinois University; Lamont, Sue (sjlamont@iastate.edu) - Iowa State University; Miller, Marcia (mamiller@coh.org) - Beckman Research Institute; Parmentier, Henk (Henk.Parmentier@wur.nl) - Wageningen Agricultural University; Parcells, Mark Parcells@udel.edu) - University of Delaware; van der Poel, Jan (jan.vanderPoel@wur.nl) - Wageningen Agricultural University; Schat, Ton (kas24@cornell.edu) - Cornell University; Scott, Tom (trscott@clemson.edu) - Clemson University; Sharif, Shayan (shayan@ovc.uoguelph.ca) - University of Guelph; Taylor, Robert (bob.taylor@unh.edu) - University of New Hamshire; Wakenell. Pat (pswakenell@ucdavis.edu)  University of California at Davis.

Administrator: Kerr, Kirklyn (Kirklyn.Kerr@uconn.edu)  University of Connecticut:

Members not present but a report has been submitted and information is included in the accomplishments: Koci, Matt (mdkoci@ncsu.edu) - North Carolina State University; Collisson, Ellen (ecollisson@westernu.edu) - Texas A & M University

Collaborator not presenting a formal report: Juul-Madsen, Helle (Helle.JuulMadsen@agrsci.dk) - Danish Institute of Agricultural Science;

Visitors: van Ginkel, Frits (vangifw@vetmed.auburn.edu) - Auburn University; Minozzi, Giulietta (giulietta.minozzi@jouy.inra.fr)  INRA, France; Pevzner, Igal (Igal.Pevzner@cobb-vantress.com)- - Arkansas; Rath, Narayan (nrath@uark.edu) - USDA, Arkansas; Rodiriguez-Lecompte, Juan Carlos (JC_Rodriguez-Lecompte@umanitoba.ca)  University of Manitoba.

Brief Summary of Minutes

Minutes of Annual Meeting
Recorded by Pat Wakenell, 2007 secretary

NE1016 business meeting opened 4:15 PM
Present: Pat Wakenell, Sue Lamont , Tom Scott, Henk Parmentier, Jan van der Poel, Renee Kopulos, Bob Taylor, Mohammad Heidari, Gisela Erf, Shayan Sharif, Fritz van Ginkel, Marcia Miller, Ton Schat, Mark Parcells, Narayan Rath, Sandy Ewald, Kirk Kerr

Kirk Kerr: explained requirement for renewal of grant every 5 years  old is closed out and new begun  time line was handed out in hard copy format.
Request for renewal must be submitted by 10/31/07, proposal must be submitted by 2/29/08 AFTER peer-review by at least 3 reviewers  group can suggest the reviewers  get to peer reviewers by 2/1/08.
From todays meeting, need minutes, annual report and a person to do the summary report of end of project (due 90 days from 10/1/08). Pat responsible for minutes and Ton responsible for annual report.
There was no annual report from last year  Bob will take care of that.
Some of new members did not submit proper forms for membership  Huaijun Zhou - Bob will set up a secondary list of invited but not formal members and ask them to submit the required forms if they still plan to be members.
No budget information available with absence of Muquarrab Qureshi.
Kirk plans to work on revision but will be stepping down July 1.
The technical committee commends Kirk for the years of service to the NE1016 committee. Kirk asked if there are any suggestions of a person in the Northeast that would replace him. Bob is in charge of the renewal of the true NE-60 / NE1016 project. Bob had emailed two requests prior to this meeting with an incredible dearth of responses. He will send 1 reminder 1 week later. After that, you will be on the tardy list that will be distributed to all members for maximum effect.
Changes of objectives: New suggestions:
Objective 1. Identify and characterize genes and their relationships to disease resistance in poultry with an emphasis on the MHC as well as other genes encoding alloantigens, communication molecules and their receptors and other candidate systems.
Objective 2. Identify and characterize environmental, dietary and physiologic, factors that modulate immune system development, optimal immune function and immune system related disease resistance and welfare in poultry genetic stocks.
Objective 3. Develop, evaluate and characterize methodologies, reagents and genotypes to assess immune function and disease resistance to enhance production efficiency through genetic selection in poultry
Publications lists needed by Fri  from 2003 no in press, format of Poultry Science, list only # of abstracts, no Endnote formats!
Based on above new objectives, need a max 2000 limit total of which objective and what will be done in new project by 11/1/07.

Accomplishments

Extensive collaboration among members continues as evidenced by the frequent exchange of reagents, birds and virus strains. Studies continued on the identification of candidate genes influencing resistance to pathogens, fine mapping of the MHC region, and on understanding vaccine induced immunity.<br /> Auburn University (AL). Broiler flocks are being established that segregate for the MX gene Asn631 and Ser631. The MX gene with Asn631 has been linked with antiviral activity against avian influenza virus (AIV) after activation by type 1 interferon (IFN-I). Chicken embryo kidney (CEK) cells were treated with IFN-± and challenged with low pathogenic (LP) AIV. Thus far few results have been analyzed to determine the antiviral effects of MX Asn631 (MX+) with our without IFN-± treatment. There was very little antiviral affect in the Mx+/- IFN-±- compared with the Mx-/- IFN-±-treated cultures, although there may be a slight antiviral affect in MX+/- versus MX-/- cells in the absence of IFN-±. A similar approach was used to determine if MX gene activation would reduce infectious bursal disease virus (IBDV) replication in chicken embryo fibroblasts (CEF). IFN-± was a strong inhibitor of IBDV replication, but this is likely to be independent of the MX genotype. <br /> Beckman Research Institute (CA). The gene map encompassing 242 kb of the MHC region has been completed. The BG2 gene is separated from the BL and BF genes by 28 other genes, many of which are most likely to be important for immune responses. The new map also allowed locating more precisely the breakpoints for some of the recombinants which differ in the resistance to Mareks disease (MD). The epitopes were defined for BF2*13 and BF2*21. The former consisted mostly of 10 amino acids with 4 anchors (positions 2, 3, 5, and 8), while the epitopes for BF2*21 consisted mostly of 10 amino acids with anchors at position 8 and 10. The knowledge of the epitopes may facilitate the development of designer vaccines. Proteomic profiling of functional studies of embryonal natural killer (NK) cells has indicated that expression of the nonclassical MHC YF7.1 inhibits NK cell activity. In addition BF1 alleles from BF13 and B21 also inhibit NK cell activity while B2*21 promotes killing. Studies are in progress to examine the specificity of the NK receptor encoded by the Blec2 gene. Preliminary data suggest that this receptor recognizes specific nonclassical MHC-I (e.g., Y7.1) alleles. <br /> Clemson University (SC). Studies were continued on the importance of thrombocytes as a source of cytokines. Broiler chickens were fed a control chick starter feed, starter feed supplemented with corticosterone, or starter feed supplemented with corticosterone and Vitamin C. These diets did not influence the production of pro-inflammatory cytokines IL-1beta, -6 and -12 in thrombocytes. The effects of in vitro LPS treatment on thrombocytes were examined using thrombocytes from 8-12 week old white leghorn hens. LPS treatment increased significantly mRNA expression of TLR4, IL-6, and COX-2 and the release of PGE2 into the culture media. Treatment with the inhibitor PD98059 did not affect expression of IL-6 and irrespective of PD98059 treatment LPS significantly elevated the expression of IL-6. However, the LPS-stimulated expression of COX-2 and release of PGE2 into culture media were dramatically decreased by PD98059 pre-treatment followed by LPS treatment. Treatment with BMS345541 significantly inhibited LPS stimulation of IL-6 and COX-2 mRNA and had a slight effect on reduction of PGE2 release from thrombocytes into culture media. The importance of the NF-ºB pathway on the expression of IL-8 in thrombocytes was also examined. Inhibition of IKK² significantly reduced Il-8 mRNA expression. These findings suggest that thrombocytes are important cells for elaboration of inflammatory mediators which can be cytokines, prostanoids and chemokines.<br /> Cornell University (NY). The specific-pathogen-free P2a (B19B19) and N2a (B21B21) flocks are continued to be maintained and fertile eggs are available for research. The MD virus (MDV) polypeptide pp38, but not the recently described splice variants A and B, is responsible for the alteration of metabolic activity in the QTP32 (dox-inducible quail cell line QT35). Because of the difference in the activity of mitochondrial succinate dehydrogenase an up-regulation in the number of transcripts of key mitochondrial enzymes in the oxidative phosphorylation pathway was expected after Dox-induction Real-time RT-PCR (qRT-PCR) assays for Japanese quail Cytochrome C oxidase-1 (Cox-1), Cytochrome B, and ATP synthase A chain showed unexpectedly that Cox-I and ATP synthase A-chain were significantly down regulated in Dox-induced cells only expressing full length pp38. Transcription of cytochrome B was more variable but not significantly different between induced and control cells. ATPase enzyme activity in isolated mitochondria was also significantly decreased after dox-induced expression of pp38. The potential use of immune complex (Ic) vaccines consisting of chicken anemia virus (CAV) with predetermined antibody levels was examined in SPF chickens. Viremia was delayed in chickens inoculated with Ic preparations and hematocrit values were not affected with the optimal dose of antibodies. Differential qPCR analysis indicated that birds were protected against challenge.<br /> Iowa State University (IA). Seventeen highly inbred lines (50  100 generations of sib matings) of defined MHC type and a broiler line were maintained. A resource population for analyzing the host genetics of early response to Salmonella (Iowa Salmonella Response Resource Population, ISRRP) has been developed over the years by crossing outbred broiler males with females of two distinct, highly inbred lines (Leghorn and Fayoumi). The ISRRP is now maintained in advanced intercross line (AIL) form, with the F12 generation being hatched in the past year. In total 2700 SNPs were analyzed for association with Salmonella enteritidis (SE) burden levels in poultry in two F8 AILs. A total of 21 SNPs were significant at a false discovery rate of 25% for SE bacterial levels (13 associated with cecal bacterial burden and 8 with spleen bacterial burden), representing 19 new candidate genes for host response to SE with a wide variation in molecular function, including apoptosis, cell signaling, and DNA repair. ARPP-21 and MLH1 were the only genes that were associated with both spleen and cecal bacterial burden. Ten SNPs were in pathways that have been previously associated with immune response to SE bacterial challenge (toll-like receptor [TLR] signaling, apoptosis, and MAPK signaling pathways). We have conducted microarray analysis of tissue samples from the ISRRP AIL F8 generation from 1-week-old birds exposed to Salmonella at one day of age, and characterized the SE-exposed tissues as relatively resistant or susceptible to bacterial colonization based upon bacterial count. Two publications outlining the results from early (< 1d) spleen responses of three genetic lines and one on F8 cecal tissue are in preparation. qRT-PCR assays were used to investigate TLR expression in cecum and spleen of three genetic chicken lines and in two AIL (8th generation) lines after challenge with Salmonella. Genetic line effect was significant on TLR RNA expression in the spleen of Salmonella infected birds, with higher TLR2 and TLR4 expression in the Fayoumi line and higher TLR5 expression in broiler line. In Salmonella challenged birds, TLR2 expression in both cecum and spleen and TLR4 expression in spleen were significantly higher at 18 h post-inoculation compared to 2 h PI. Infection of the AIL chickens resulted in significant upregulation of TLR2 and TLR5 RNA expression in spleens and upregulation of TLR4 RNA expression in both cecum and spleen. The increase in TLR2, TLR4 and TLR5 RNA expression level was found in spleens from male but not female chicks. Bacterial burden of Salmonella serovar Enteritidis in challenged birds was not correlated with TLR RNA expression level. The impact of SE infection on lymphocytes was examined by immunohistochemistry, infection was associated significantly associated with an increased area staining for macrophages and decreased apoptosis in cecal tissue sections compared to uninfected animals. <br /> North Carolina State University (NC). Previously, the effect of the probiotic Primalac on the physiology of the gut was examined but the biological relevance of minor differences was not clear. Current studies are examining a potential influence on the immune response to sheep red blood cells (SRBC). Preliminary data suggest that Primalac may influence the kinetics and class switching of the antibody response. Type-2 turkey astrovirus (TAstV-2) is associated with the poult early mortality syndrome (PEMS), but there is no clear adaptive immune response to TAstV-2. To better understand the local immune responses to this gut pathogen qRT-PCR assays are being developed for 14 genes believed to be expressed in mucosal immunity.<br /> Northern Illinois University (IL). The RFLP patterns of Y haplotypes vary between lines to the extent that each chicken population seems to have unique Y haplotypes. Currently there are at least 30 different YF haplotypes segregating in the lines that have been analyzed. Four additional recombinant lines have been partially characterized using the LEIO258 marker. Four commercial lines, C1, C2, C3 and C6 have been tested for several of the non-MHC alloantigen systems in addition to B and Y system genotyping for progeny evaluation. These lines are currently used to produce line specific typing reagents. Proteins representing the L alloantigen have been submitted to Dr. Shane Burgess at Mississippi State University for analysis. Specific matings are conducted to supply collaborators Lillehoj and Collisson with hatching eggs for projects on coccidiosis and AIV, while Wakenell is evaluating 6 allotypes for potential resistance to MD.<br /> Texas A&M University. Tools and assays have been developed to start the evaluation of T lymphocyte responses to AIV. MHC-defined kidney cells from 10-day old naive birds will be used as a source of MHC compatible and mismatched antigen presenting cells (APC). Immune cells are collected from the peripheral blood mononuclear cells (PBMC) of AIV infected chickens 2 to 25 weeks of age and 10 days to 5 months p.i. Infected APCs and T cells are co-cultured for 24 to 48 hours prior to adding macrophage generated NO containing supernatant fluids and assayed for interferon-gamma production. Attwaters prairie chickens are a federally endangered grouse species. Captive breeding programs are hampered by the high incidence of reticuloendotheliosis virus (REV). In infected birds as much as 50% of the CD8+ or CD4+ cells in a preparation of PBMC can be actively infected with REV. Using nested PCR approaches rather than single PCR it was possible to generate a small nucleus of negative birds for breeding. However, infection could be reintroduced perhaps through poxviruses carrying REV. To prevent reinfection an experimental DNA vaccine has been developed which will be tested in the related, but not endangered, greater prairie chicken. <br /> University of Arkansas (AR). The onset of vitiligo in the Smyth line chickens is preceded and accompanied by increases in IFN-³, IL-1², IL-8 and IL-12 expression but not IL-4, iNOS and IL-6 expression based on qRT-PCR studies. These observations are in line with reports of a Th1 polarized response in human vitiligo. Moreover, NALP1, an intracellular pattern recognition receptor which detects danger signals such as microbial products and HSP and forms part of the inflammasome, is required for the cleavage of IL-1² into its active form and was recently identified as a candidate gene involved in human vitiligo and other autoimmune/immunoinflammatory diseases. The expression of NALP1 in feathers from SL chickens with vitiligo is increased 3-fold above levels observed in feathers from SL chickens without vitiligo and parallels the increase in IL-1². Studies are in progress to determine the potential involvement of macrophages, complement and autoantibodies in vitiligo. The latter may be used as indicators of antigens involved in the melanocyte destruction. A time course study on the role of herpesvirus of turkeys (HVT) in vitiligo is in progress. Feather tips were found to be a good source for the study of immune cells and antibody responses. Antigen can be injected in the feather tips and the tips as well as blood samples can than be used to monitor local as well as systemic immune responses.<br /> University of California-Davis (CA). The resistance to MD of an additional recombinant, BR3, was compared to the MD resistance of BR2 and BR4. The percentage of MD positive BR3 homozygous birds (12) was intermediate between that of the BR2 homozygotes (8) and the BR4 homozygotes (16), but histopathology is still pending. The pathogenicity of QMDV, a MDV isolate rescued from QT35 cells, was compared with JM, RB1B, and NC584A (chickens only) in both SPF chickens and quail. Preliminary results indicate that QMDV is comparable to RB1B in chickens and JM in quail. Serum chemistry was performed on SPF chickens challenged with Eimeria tenella. Serum AP, LK aspartamine aminotransferease (AST), GGT and LDH were significantly lower in E. tenella challenged birds at day 5 PC then in control birds. When serum chemistries were compared between birds which died during this period and those that survived, AP and AST were significantly depressed in moribund birds at day 5 post infection but there were no differences with CK, GGT and LDH. <br /> University of Delaware (DE). The MDV Meq gene has been linked to transformation, but may have also other functions. Meq genes in lower virulence MDVs (notably JM102, CU-2, BC-1, etc.) have an expansion of the proline-rich repeat region, which is associated with transcriptional repression. In vv+MDVs Meq has specific mutations in this proline rich repeat region with functional consequences for the transcriptional activation function of this protein. Research is focused on the identification of actin-regulating genes as a class of Meq target genes and to determine if Meq-induced changes can influence the susceptibility of tumor target cells to cell-mediated immune responses. <br /> The importance of MDV glycoprotein L (gL) was examined by using a gL deletion mutant (TK-1a) of the TK strain. Antibodies to gL, gH and the gH/gL complex (generated against MDV-1, 584c p80, or CVI988 strains) cross-react strongly with gL and gH of HVT and SB1. These data suggest that not only did the vaccines present similar epitopes to the gL/gH complex, but that HVT expresses a higher-density of these antigens per infected cell than MDV-1 strains. The gL deletion was observed in all field isolates of MDV-1 (n=30) obtained from vaccine breaks in 2005 and 2007. To examine the imporetance of this deletion for vaccine-induced protection, commercial broiler embryonated eggs were vaccinated in ovo at ED 18 with HVT (5,000 PFU), with bivalent vaccine (5,000 PFU HVT/ 2,500 PFU SB1) or left unvaccinated and inoculated with 200 PFU of TK-1a (gL ), TK-2a (gL non-deleted) or a 1:1 mix of these strains (Mix) at hatch. The incidence of MD + Mortality was decreased by vaccination (HVT and HVT/SB1 groups) indicating that even with challenge at day of age, vaccination provides a level of protection. Despite having a decreased incidence of MD + Mortality in vaccinated groups, the TK-1a strain caused high levels of MD + Mortality in both HVT and HVT/SB1 vaccinated groups. Challenge with TK2a showed the expected pattern of incidence with HVT providing some protection, but the bivalent vaccination provided higher levels of protection. Birds infected with theTK-2a and the Mix of TK-1a +TK-2a experienced a greater level of MD + Mortality than the TK-1a strain alone. Analysis of virus in the birds inoculated with Mix showed predominantly TK-1a virus.<br /> University of Guelph (CANADA). Immune-specific Microarrays have been used to compare immune system gene responses in the Cornell P2a and N2a chickens at 4-21 days post infection with the RB1B strain. Genes associated with T and B cell and perhaps NK cells were differentially expressed during this period. T cell subsets from RB1B-infected P2a and N2a chickens were prepared at 4, 10 and 21 days post infection and analyzed for expression of cytokine mRNA by qRT-PCR. IL-18 was upregulated at 4 days in CD4+ cells and at 21 days in CD8+ cells in the susceptible line and at 4 and 21 days in CD8+ cells in the resistant line. Il-6 was upregulated at 4 days in both lines, while IL-10 was expressed at all time points in both lines. CD4+ and CD8+ cells infiltrate in the feather pulp of MDV-infected chickens starting at 4 days post infection with peak infiltration at 10 days. The expression of IL-18, IL-6, interferon-gamma and MHC-I was significantly enhanced in the feather pulp of MDV-infected chickens.<br /> University of New Hampshire (NH). Six-week-old R13R13 chickens were compared to other MHC congenic recombinant (R1, R2, R4, R5) lines for Rous sarcoma virus (RSV) tumor outcome after challenge with subgroup C RSV. Tumors grew in >90% of inoculated chickens. The tumor size increased in all genotypes through week 3 post-inoculation. Tumors in R2R2, R4R4, and R5R5 chickens plateaued thereafter while tumors in R1R1 chickens continued to increase in size. The mean tumor profile index (TPI) for R5R5 chickens was significantly lower than the TPI of R2R2, R4R4, and R13R13, but not R1R1.<br /> USDA, Avian Disease and Oncology Laboratory. A formal report was not presented due to the change in personnel associated with the NE-1016 project.<br /> Wagenigen Agricultural University (NL). The 20K SNP set (Illumina Infinium platform) are being used to genotype a broiler chicken population and several research chicken lines from the IAH (Compton, UK), INRA and the Wageningen selection lines (high, low and control). The sequences of the first domain of the B-F locus and the BLb2 locus were analyzed in the Wageningen lines showing variation. Thus, sequencing of the BF1 or BLb2 gene exon seems a necessary addition to the molecular MHC typing using the LEI0258 microsatellite marker. Based on the DNA sequence analysis two B24 subtypes could be defined in animals, which seemed homozygous using the LEI0258 marker. One of the Wageningen B24 subtypes is identical to B24 defined by Briles, the other is clearly distinct. LEI0258 homozygous chickens from the INRA lines also showed polymorphism in TAP1, TAP2, tapasin, BF and BL genes. A SNP set representing 1536 SNP focusing on the QTL region on GGA 3, 4, 5, 7, 26 and Z has been used to genotype 960 animals, the resulting genotypes are being are being used for associations studies with previously recorded immunological and behavioral parameters. PAMP-like components present in organic feed may modulate immune responses. The proposed working hypothesis is that organic feed may restore immune functions to a normal species-restricted average level. To determine if alpha-GAL has an adjuvant function, layers were inoculated with alpha-GAL Human serum albumin (HuSA), beta-GAL HuSA, or naïve HuSA. Unexpectedly, the addition of alpha-GAL, but not beta-GAL, decreased antibody responses to HuSA significantly. In preliminary studies the addition of dietary probiotics did not influence immune competence in layers. Studies were initiated to identify physiological (behavioral, immunological) mechanisms and parameters that predict and/or underlie health status and zootechnical characteristics but also abnormalities (feather pecking, cannibalism, and unidentified death causes) in various layer breeds kept under commercial conditions. The first results suggest a relation between level of binding of natural antibodies to LPS or KLH and risk of mortality. Additional data suggest different levels of (heat-) stress sensitivity in layer breeds which were established at the neuron-endocrine as well as the immunological level, and which were reflected at the growth and production level. The data suggest that birds that are able to mount higher corticosteron responses to stress are more able to respond and/or adapt to stress than birds with lower corticosteron responses.<br />

Publications

Impact Statements

1. The fine mapping of the MHC region will facilitate a better understanding of the differences between birds in immune responses to vaccines and infections.
2. The major effort in understanding the genetic basis for differences in innate and acquired immunity will benefit the industry in providing tools for selection.
3. Genetic resources are still maintained at several institutes, but the continued financial pressures may decrease the availability of important resources in the future.

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Participants

Bob Taylor, Sue Lamont, Kirk Klasing, Sandra Ewald, Shayan Sharif, Mary Delany, Pat Wakenell, Tom Scott, Muquarrab Qureshi, Jan van der Poel, Henk Parmentier, Matt Koci

Brief Summary of Minutes

Accomplishments

Publications

Impact Statements

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Report Information

Participants

Collisson, Ellen (ecollisson@westernu.edu) - Western University;

Delany, Mary (medelany@ucdavis.edu) - University of California at Davis;

Erf, Gisela (gferf@uark.edu)- University of Arkansas;

Ewald, Sandra (ewaldsj@auburn.edu) - Auburn University;

Klasing, Kirk (kcklasing@ucdavis.edu) - University of California at Davis;

Koci, Matt (mdkoci@ncsu.edu)- North Carolina State University;

Lamont, Sue (sjlamont@iastate.edu) - Iowa State University;

Miller, Marcia (mmiller@coh.org) - City of Hope;

Parcells, Mark (parcels@udel.edu) - University of Delaware;

Schat, K. A. (kas24@cornell.edu) - Cornell University;

Scott, Tom (trscott@clemson.edu) - Clemson University;

Sharif, Shayan (shayan@ovc.uoguelph.ca) - University of Guelph;

Taylor, Bob (bob.taylor@unh.edu) - University of New Hampshire;

Wakenell, Pat (pwakenel@purdue.edu) - Purdue University

Brief Summary of Minutes

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

Accomplishments

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.<br /> <p><br /> 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.<br /> <p><br /> 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.<br /> <p><br /> 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.<br /> <p><br /> 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. <br /> <p><br /> 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.<br /> <p><br /> 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.<br /> <p><br /> 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. <br /> 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.<br /> <p><br /> 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.<br /> <p><br /> 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.<br /> <p><br /> 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.<br /> 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. <br /> <p><br /> 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.<br /> <p><br /> 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.<br /> 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.<br /> <p><br /> 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).<br /> <p><br /> 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.<br /> <p><br /> 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.<br /> <p><br /> 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.<br /> 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.<br /> <p><br /> 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.<br /> <p><br /> 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.<br /> <p><br /> 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.<br /> <p><br /> 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.<br /> <p><br /> 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.<br /> <p><br /> 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.<br /> <p><br /> 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.<br /> <p><br /> 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. <br /> <p><br /> 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.<br /> <p><br /> Objective 3: Develop and evaluate methodologies and reagents to assess immune function and disease resistance to enhance production efficiency through genetic selection in poultry.<br /> <p><br /> 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. <br /> <p><br /> 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.<br /> <p><br /> 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. <br /> <p><br /> 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.<br /> <p><br /> 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.<br /> <p><br /> 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.<br /> 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.<br /> <p><br /> 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.<br /> 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.<br /> <p><br /> 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.<br />

Publications

Abasht, B., J. C. M. Dekkers, and S. J. Lamont. 2006. Review of quantitative trait loci identified in the chicken. Poult. Sci. 85:2079-2096.<br /> <br /> Abasht, B., M. G. Kaiser, and S. J. Lamont. 2008. Toll-like receptor gene expression in cecum and spleen of advanced intercross line chicks infected with Salmonella enterica serovar Enteritidis, Vet. Immuno. Immunopathol. 123:314-323.<br /> <br /> Abdul-Careem, M. F., B. D. Hunter, É. Nagy, L. R. Read, B. Sanei, J. L. Spencer, and S. Sharif. 2006. Development of a real-time PCR assay using SYBR Green chemistry for monitoring Marek's Disease Virus genome load in feather tips. J. Virol. Methods 133:34-40.<br /> <br /> Abdul-Careem, M. F., B. D. Hunter, A. J. Sarson, A. Mayameei, H. Zhou, and S. Sharif. 2006. Marek's disease virus-induced transient paralysis is associated with cytokine gene expression in the nervous system. Viral Immunol. 19:167-176.<br /> <br /> Abdul-Careem, M. F., B. D. Hunter, P. Parvizi, H. R. Haghighi, N. Thanthrige-Don, and S. Sharif. 2007. Cytokine gene expression patterns associated with immunization against Marek's disease in chickens. Vaccine 25:424-432.<br /> <br /> Abdul-Careem, M. F., D. B. Hunter, M. D. Lambourne, J. Barta, and S. Sharif. 2007. Ontogeny of cytokine gene expression in the chicken spleen. Poult. Sci. 86:1351-1355.<br /> <br /> Abdul-Careem, M. F., D. B. Hunter, M. D. Lambourne, L. R. Read, P. Parvizi, and S. Sharif. 2008. Expression of cytokine genes following pre- and post-hatch immunization of chickens with herpesvirus of turkeys. Vaccine 26:2369-77.<br /> <br /> Abdul-Careem, M. F., D. B. Hunter, N. Thanthrige-Don, H. R. Haghighi, M. D. Lambourne, S. Sharif. 2008. Cellular and cytokine responses associated with dinitrofluorobenzene-induced contact hypersensitivity in the chicken. Vet. Immunol. Immunopath. 122:275-84.<br /> <br /> Abdul-Careem, M. F., B. D. Hunter, A. J. Sarson, P. Parvizi, H. R. Haghighi, L. Read, M. Heidari, and S. Sharif . 2008. Host responses are induced in feathers of chickens infected with Marek's disease virus. Virol. 370:323-32.<br /> <br /> Anobile, J. M., V. Arumugaswami, D. Downs, K. Czymmek, M. Parcells, and C. J. Schmidt. 2006. Nuclear Localization and Dynamic Properties of the Marek's Disease Virus Oncogene Products Meq and Meq/vIL8. J. Virol. 80:1160-1166.<br /> <br /> Ardia, D. R., and K. A. Schat. 2008. Ecoimmunology. Pages 421-442 in Avian Immunology. F. Davison, B. Kaspers, and K. A. Schat, eds. Elsevier Academic Press, London.<br /> <br /> Asif, M., J. W. Lowenthal, M. E. Ford, K. A. Schat, W. G. Kimpton, and A. G. D. Bean. 2007. IL-6 expression following IBV infection in chickens. Viral Immunol. 20:479-486.<br /> <br /> Blackmore, C., K. Klasing, and P. Wakenell. 2006. Effect of infectious bursal disease virus insult on iron, copper, and zinc concentration in liver, bursa of fabricius, spleen, pancreas, and serum of chickens. Avian Dis. 50:303-305.<br /> <br /> Blatchford, R. A., K. C. Klasing, H. L. Shivaprasad, P. S. Wakenell, G. S. Archer, and J. A. Mench. 2009. The effect of light intensity on the behavior, eye and leg health, and immune function of broiler chickens. Poult. Sci. 88:20-28.<br /> <br /> Bowen, O. T., G. F. Erf, N. B. Anthony, and R. F. Wideman. 2006. Pulmonary hypertension triggered by lipopolysaccharide (LPS) in ascites-susceptible and resistant broilers is not amplified by aminoguanidine, a specific inhibitor of inducible nitric oxide synthase (iNOS). Poult. Sci. 85:528-536.<br /> <br /> Bowen, O. T., R. F. Wideman, N. B. Anthony, and G. F. Erf. 2006. Is variation in the pulmonary hypertensive responsiveness of broilers to lipopolysaccharide (LPS) attributable to innate variation in nitric oxide (NO) production by mononuclear cells? Poult. Sci. 85:1349-1363.<br /> <br /> Bowen, O. T., G. F. Erf, M. E. Chapman, and R. F. Wideman, Jr. 2007. Plasma nitric oxide concentration in broilers after intravenous injections of lipopolysaccharide or microparticles. Poult. Sci. 86:2550-54.<br /> <br /> Brentano, L., S. Lazzarin, T. A. P. Klein, and K. A. Schat. 2005. Detection of chicken anemia virus in the gonads and in the progeny of broiler breeder hens with high neutralizing antibody titer. Vet. Microbiol. 105:65-72.<br /> <br /> Brisbin, J. T., H. Zhou, J. Gong, P. Sabour, M. R. Akbari, H. R. Haghighi, H. Yu, A. Clarke, A. J. Sarson, and S. Sharif (2008) Gene expression profiling of chicken lymphoid cells after treatment with Lactobacillus acidophilus cellular components. Dev. Comp. Immunol. 32:563-74.<br /> <br /> Brown, D. C., C. V. Maxwell, G. F. Erf, M. E. Davis, S. Singh, and Z. B. Johnson. 2006. The influence of different management systems and age on intestinal morphology, immune cell numbers and mucin production from goblet cells in post-weaning pigs. Vet. Immunol. Immunopathol. 111:187-98.<br /> <br /> Burgess, S. C., J. R. Young, B. J. Baaten, L. Hunt, L. N. Ross, M. S. Parcells, P. M. Kumar, C. A. Tregaskes, L. F. Lee, and T. F. Davison. 2004. Marek's disease is a natural model for lymphomas overexpressing Hodgkin's disease antigen (CD30). Proc Natl. Acad. Sci. USA 101:13879-13884.<br /> <br /> Buyse, J., Q. Swennen, T. A. Niewold, K. C. Klasing, G. P. Janssens, M. Baumgartner, and B. M. Goddeeris. 2007. Dietary l-carnitine supplementation enhances the lipopolysaccharide-induced acute phase protein response in broiler chickens. Vet. Immunol. Immunopathol. 118: 154-159.<br /> <br /> Buyse, J., Q. Swennen, F. Vandemaele, K. C. Klasing, T. A. Niewold, M. Baumgartner, and B. M. Goddeeris. 2008. Dietary beta-hydroxy-beta-methylbutyrate supplementation influences performance differently after immunization in broiler chickens. J. Anim. Physiol. Anim. Nutr. 26:11-19.<br /> <br /> Chapman, M. E., W. Wang, G. F. Erf, and R. F. Wideman, Jr. 2005. Pulmonary hypertensive responses of broilers to bacterial lipopolysaccharide (LPS): evaluation of LPS source and dose, and impact of pre-existing pulmonary hypertension and cellulose microparticle selection. Poult. Sci. 84:432-441.<br /> <br /> Chapman, M. E., R. L. Taylor, Jr., and R. F. Wideman, Jr. 2008. Analysis of plasma serotonin levels and hemodynamic responses following chronic serotonin infusion in broilers challenged with bacterial lipopolysaccharide and microparticles. Poult. Sci. 87:116-124.<br /> <br /> Cheeseman, J. H., M. G. Kaiser, C. Ciraci, P. Kaiser, and S. J. Lamont. 2007. Breed effect on early cytokine mRNA expression in spleen and cecum of chickens with and without Salmonella enteritidis infection. Dev. Comp. Immunol. 31:52-60. <br /> <br /> Cheeseman, J. H., M. G. Kaiser, and S. J. Lamont. 2004. Genetic line effect on peripheral blood leukocyte cell surface marker expression in chickens. Poult. Sci. 83:911-916. <br /> <br /> Cheeseman, J. H., N. A. Levy, P. Kaiser, H. S. Lillehoj, and S. J. Lamont. 2008. Salmonella enterica serovar Enteritidis induced alteration of inflammatory CXCL chemokines mRNA expression and histological changes in the cecum of infected chicks. Avian Dis. 52:229-234. <br /> <br /> Cheeseman, J. H., H. S. Lillehoj, and S. J. Lamont. 2008. Reduced nitric oxide production and iNOS mRNA expression in IFN-g-stimulated chicken macrophages transfected with iNOS siRNAs. Vet. Immunol. Immunopathol. 125:375-380. <br /> <br /> Cheng, H. H., and S. J. Lamont. 2008. Genetics of disease resistance. Pp. 59-72. In: Diseases of Poultry. 12th ed. Y.M. Saif, A. Fadly, J. Glisson, I. McDonald, L. Nolan, and D. Swayne, Eds., Blackwell Publ.<br /> <br /> Cheng, H., M. Niikura, T. Kim, W. Mao, K. S. MacLean, H. Hunt, J. Dodgson, B. Burnside, R. Morgan, M. Ouyang, S. Lamont, J. Dekkers, J. Fulton, M. Soller, and W. Muir. 2008. Using integrative genomics to elucidate genetic resistance to Marek's disease in chicken. In : Pinard M-H, Gay C, Pastoret P-P, Dodet B (eds): Animal Genomics for Animal Health. Dev Biol (Basel). Basel, Karger, 2008, vol 132, p 365-372.<br /> <br /> Chichlowski, M., J. Croom, B. W. McBride, L. R. Daniel, G. Davis and M. D. Koci. Direct-Fed Microbial and Salinomycin Modulate Whole Body and Intestinal Oxygen Consumption and Intestinal Mucosal Cytokine Production in the Broiler Chick. Poult. Sci. 86:1100-1106, 2007.<br /> <br /> Chichlowski, M., J. Croom, F. Edens, B. W. McBride, R. Qiu, C. C. Chiang, L. R. Daniel, G. W. Havenstein and M. D. Koci. 2007. Microarchitecture and Spatial Relationship Between Bacteria and Ileal, Cecal and Colonic Epithelium in Chicks Fed a Direct-Fed Microbial and Salinomycin. Poult. Sci. 86:1121-1132. <br /> <br /> Chichlowski, M., J. Croom, B.W. McBride, G. B. Havenstein and M. D. Koci. 2007. Metabolic and Physiological Impact of Probiotics or Direct-Fed-Microbials on Poultry: A Brief Review of Current Knowledge. Int. J. Poult. Sci. 6:694-704.<br /> <br /> Collisson, E. W., S. Singh, and T. Omran, 2007. Developments in avian influenza virus vaccines. J Poult. Sci. 44 (No.3):238-257.<br /> <br /> Collisson, E. W., S. Singh, and Y. Drechsler. 2008. Evolving vaccine strategies for the continuously evolving avian influenza viruses. CAB Rev: Perspectives in Vet. Med., Agric. Nutr. and Nat. Resources.<br /> <br /> Cumberbatch, J.A., D. Brewer, I. Vidavsky, and S. Sharif. 2006. Chicken major histocompatibility complex class II molecules of B19 haplotype present self and foreign peptides. Anim. Genet. 37:393-396.<br /> <br /> Davison, F., B. Kaspers, and K.A. Schat, eds. 2008. Avian Immunology. F. Davison, B. Kaspers, and K.A. Schat, eds. Elsevier Academic Press, London.<br /> <br /> Davison, F., B. Kaspers, and K. A. Schat. 2008. The avian mucosal immune system. Page 241-242 in Avian Immunology. F. Davison, B. Kaspers, and K. A. Schat, eds. Elsevier Academic Press, London.<br /> <br /> Erf, G. F. 2004. Avian cell-mediated immunity. Poult. Sci. 83:580-590.<br /> <br /> Erf, G. F. 2008. Autoimmune diseases of poultry. Pages 339-358 in: Avian Immunology, Davison F., Kaspers, B., and K. A. Schat, editors. Elsevier, London, San Diego, CA.<br /> ,<br /> Erf, G. F., and R. L. Taylor, Jr. 2004. Introduction: Ancillary Scientist Symposium on the avian immune system - function and modulation. Poult. Sci. 83:580.<br /> <br /> Ewald, S. J., and E. J. Livant. 2004. Distinctive polymorphism of chicken B-FI (MHC class I) molecules. Poult. Sci. 83:600-645.<br /> <br /> Ewald, S. J., X. Ye, S. Avendano, S. McLeod, S. J. Lamont, and J. C. Dekkers. 2007. Associations of BF2 alleles with antibody titres and production traits in commercial pure line broiler chickens. Anim. Genet. 38:174-176.<br /> <br /> Fan, Y.K., W. J. Croom, Jr., Linda Daniel, B.W. McBride, M. Koci, G. B. Havenstein , and E. J. Eisen. 2006. Effect of Genotype on Whole-Body and Intestinal Metabolic Response to Monensin in Mice. Asian-Australasian J. Anim. Sci. 19:554-562.<br /> <br /> Ferdous, F., D. V. Maurice, and T. R. Scott. 2008. Broiler chick thrombocyte response to lipopolysaccharide. Poult. Sci. 87:61-63.<br /> <br /> Fulton, J. E., H. Juul-Madsen, C. M. Ashwell, A. M. McCarron, J. A. Arthur, N. O'Sullivan, and R. L. Taylor, Jr. 2006. Molecular genotype identification of the Gallus gallus major histocompatibility complex. Immunogenetics 58:407-421.<br /> <br /> Garcia Camacho, L., K. A. Schat, R. Brooks, Jr., and D. I. Bounous. 2003. Early cell-mediated immune responses to Marek's disease virus in two chicken lines with defined major histocompatibility complex antigens. Vet. Immunol. Immunopathol. 95:145-153.<br /> <br /> Garcia-Espinosa, G., S. Clerens, L. Arckens, G. F. Erf, G. Tellez, and B.M. Hargis. 2008. Peptides from the bursa of Fabricius associated with suppression of mitogen stimulated DNA-synthesis in burs of Fabricius cells belong to intracellular proteins. Int. J. Poult. Sci. 7:125-128.<br /> <br /> Ghebremichael, S. B., J. R. Hasenstein, and S. J. Lamont. 2008. Association of interleukin-10 cluster genes and Salmonella response in the chicken. Poult. Sci. 87:22-26. <br /> <br /> Goodenbour, J. M., M. G. Kaiser, and S. J. Lamont. 2004. Linkage mapping of inhibitor of apoptosis protein-1 (IAP 1) to chicken chromosome 1. Anim. Genet. 35:158-159.<br /> <br /> Haeri, M., L. R. Read, B. N. Wilkie, and S. Sharif. 2005. Identification of peptides associated with chicken major histocompatibility complex class II molecules of B21 and B19 haplotypes. Immunogenetics 56:854-859.<br /> <br /> Haghighi, H. R., J. Gong, C. L. Gyles, M. A. Hayes, B. Sanei, P. Parvizi, H. Gisavi, J. R. Chambers, and S. Sharif. 2005. Modulation of antibody-mediated immune response by probiotics in chickens. Clin. Diagn. Lab. Immunol. 12:1387-1392.<br /> <br /> Haghighi, H. R., J. Gong, C. L. Gyles, M. A. Hayes, H. Zhou, B. Sanei, J. R. Chambers, and S. Sharif. 2006. Probiotics stimulate the production of natural antibodies in chickens. Clin. Vaccine Immunol. 13:975-980.<br /> <br /> Haghighi, H. R., M. F. Abdul-Careem, R. A. Dara, J. R. Chambers, and S. Sharif. 2008. Cytokine gene expression in chicken cecal tonsils following treatment with probiotics and Salmonella infection. Vet. Microbiol. 126:225-233.<br /> <br /> Hamal, K., S. Burgess, I. Pevzner, and G. F. Erf. 2006. Maternal antibody transfer from dams to their egg-yolk, egg-white and chicks in two meat lines of chickens. Poult. Sci. 85:1364-1372.<br /> <br /> Hamal K. R., R. F. Wideman, Jr., N. B. Anthony, and G. F. Erf. 2008. Expression of inducible nitric oxide synthase in lungs of broiler chickens following intravenous cellulose microparticle injection. Poult. Sci. 87:636-44.<br /> <br /> Hangalapura, B. N., M. G. Kaiser, J. J. Poel, H. K. Parmentier, and S. J. Lamont. 2006. Cold stress equally enhances in vivo pro-inflammatory cytokine gene expression in chicken lines divergently selected for antibody responses. Dev. Comp. Immunol. 30:503-511. <br /> <br /> Hasenstein, J. R., A. T. Hassen, J. C. M. Dekkers, and S. J. Lamont. 2008. High resolution, advanced intercross mapping of host resistance to Salmonella colonization. In : Pinard M-H, Gay C, Pastoret P-P, Dodet B (eds): Animal Genomics for Animal Health. Dev. Biol. (Basel). Basel, Karger, 2008, vol 132, p 213-218.<br /> <br /> Hasenstein, J. R., and S. J. Lamont. 2007. Chicken gallinacin gene cluster associated with Salmonella response in advanced intercross line. Avian Dis. 51:561-567. <br /> <br /> Hasenstein, J. R., G. Zhang, and S. J. Lamont. 2006. Analyses of five gallinacin genes and the Salmonella enterica serovar Enteritidis response in poultry. Infect. Immun. 74:3375-3380.<br /> <br /> Heidari, M., H. M. Zhang, and S. Sharif. 2008. Marek's disease virus induces Th2 activities during the cytolytic infection. Viral Immunol. 21:203-14.<br /> <br /> Herlong, J. L., and T. R. Scott. 2006. Positioning prostanoids of the D and J series in the immunopathogenic scheme. Immunol. Let. 102:121-131.<br /> <br /> Higgins, S. E., G. F. Erf, J. P. Higgins, S. N. Henderson, A. D. Wolfenden, G. Gaona-Ramirez, and B. M. Hargis. 2007. Effect of probiotic treatment in broiler chicks on intestinal macrophage numbers and phagocytosis of Salmonella enteritidis by abdominal exudate cells. Poult. Sci. 86:2315-21.<br /> <br /> Hillier, L. W., W. Miller, E. Birney, W. Warren, R. C. Hardison, C. P. Ponting, P. Bork, D. W. Burt, M. A. Groenen, M. E. Delany, J. B. Dodgson, A. T. Chinwalla, P. F. Cliften, S. W. Clifton, K. D. Delehaunty, C. Fronick, R. S. Fulton, T. A. Graves, C. Kremitzki, D. Layman, V. Magrini, J. D. McPherson, T. L. Miner, P. Minx, W. E. Nash, M. N. Nhan, J. O. Nelson, L. G. Oddy, C. S. Pohl, J. Randall-Maher, S. M. 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Impact Statements

1. Progress was made toward a greater understanding of the roles of the chicken major histocompatibility complex (MHC) genes in disease resistance and immune potential. Translation of knowledge in this area has direct impact on the selection schemes used by poultry industry breeders for the production of more disease resistant lines of chickens.
2. Utilization of genetic background information, particularly MHC, has allowed for evaluation of poultry performance under various environmental conditions including diseases. Knowledge of genetic backgrounds provides for development of strategies in the field to overcome stressors and unusual situations that can challenge the performance of chickens.
3. In concert with knowledge gained about the genetics of poultry, improved methods for evaluation of immune responsiveness and disease resistance have been incorporated into assessments of performance. A wider range of understanding about poultrys use of innate and adaptive modes of immunity have provided for improved and sensitive methods of performance evaluation for health status.

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