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use of mouse models has provided valuable insight into the etiology of monogenic syndromes caused by single gene mutations. However, such models do not mimic the genetic complexity of disease traits commonly seen in the human population. Complex traits, such as polygenic growth and obesity are influenced by the small to moderate direct effects of quantitative trait loci (QTL), epistasis between QTL alleles, environmental perturbations and QTL-environment interactions.\nTo date numerous mouse growth and obesity QTL have been localized [1,2], however, little progress has been made in determining the specific nature of the underlying genetic variants. One resource used to fine map QTL are congenic strains which are designed to convert a complex polygenic trait into one that is mono- or oligogenic. This is accomplished by eliminating segregating genetic variation outside the locus of interest and reducing the environmental variation influencing a trait by characterizing large numbers of genetically identical mice. The Complex Trait Consortium (CTC) considers congenic analysis an excellent method to confirm and subsequently fine map QTL [3].\nTraditionally, congenic strains are developed by introgressing QTL alleles from a donor strain, whose boundaries are defined by genetic markers, on the genetic background of a recipient strain via 10 backcrosses [4]. While technically straightforward this is a time consuming endeavor taking over three years to construct a single strain. The speed congenic approach is an alternative to this lengthy process and can reduce the number of required backcrosses from 10 to five [5,6]. This strategy uses marker-assisted selection to identify male mice inheriting fewer donor alleles, than expected on average, during each backcross. Numerous traditionally developed and speed congenics have been used to successfully isolate mouse QTL for a wide array of traits, including growth and obesity [7-13].\nThe C57Bl/6J-hg/hg (HG) mouse is a model of systemic overgrowth resulting from a spontaneous deletion on MMU10, which eliminates expression of the Socs2 (suppressor of cytokine signaling 2) gene [14-16]. The role of Socs2 in the HG phenotype was confirmed by an independently engineered Socs2-/- knockout mouse which shared a number of phenotypes in common with HG, including gigantism [17].\nQTL which alter the phenotypic effects of another locus are referred to as modifier QTL [18]. Modifier QTL have been mapped for numerous traits and in these studies the modified locus is typically a known gene containing a spontaneously arisen or engineered mutation with major phenotypic effects, such as hg [11,19-21]. Epistasis forms the basis of these interactions, implying that the known gene and its modifiers are members of the same biochemical or metabolic pathway. Different genetic backgrounds have been shown to modify the growth-enhancing effects of hg [15,22]. Since the primary function of Socs2 is to negatively regulate growth hormone (Gh) [23], it is likely these background effects are the result of polymorphism influencing interactions between members of the Gh signaling pathway. Thus, identification of hg modifiers has the potential to uncover novel members of metabolically important pathways or previously unknown convergences between pathways.\nAs an initial step, growth and carcass composition QTL were identified in a cross between CAST/EiJ (CAST) and HG [24]. In the F2, mice homozygous for the hg deletion (hg/hg) and wild type (+/+) were assayed and hg modifier QTL were defined as those either absent in +/+ mice and segregating in hg/hg or loci with altered gene action dependent on background. Four hg modifier QTL (Wg2 on MMU2, Carfhg2 on MMU9, Carp2 on MMU11 and Feml3 on MMU17) were identified, along with 12 additional QTL (Q1Ucd1 on MMU1 (which only reached a suggestive level of statistical significance), Wg1, Carp1, Cara1 and Feml1 on MMU2, Carfhg1 on MMU5, Wg3 on MMU8, Feml2 on MMU9, Wg4 and Cara2 on MMU11 and Carp3 and Cara3 on MMU17) whose expression was independent of the hg locus.\nIn the current study we have developed speed congenic strains to isolate all the aforementioned QTL. Single strains on an HG background were created for each chromosomal region outside of MMU2, while a comprehensive panel of overlapping strains with identical donor regions on both B6 and HG backgrounds were developed for MMU2. The MMU2 panel was developed to confirm the presence of multiple QTL and test for QTL-hg interactions. Additionally, based on the knowledge of potential QTL-hg interactions, genes involved in Gh signaling, whose genomic location overlapped hg modifier QTL on MMU2, 9, 11 and 17, were sequenced."}

{"project":"craft-ca-core-ex-dev","denotations":[{"id":"T2618","span":{"begin":22,"end":27},"obj":"NCBITaxon:10088"},{"id":"T2619","span":{"begin":90,"end":95},"obj":"SO_EXT:0000704"},{"id":"T2620","span":{"begin":123,"end":127},"obj":"SO_EXT:0000704"},{"id":"T2621","span":{"begin":128,"end":137},"obj":"SO_EXT:sequence_alteration_entity_or_process"},{"id":"T2622","span":{"begin":177,"end":184},"obj":"SO_EXT:0000704"},{"id":"T2623","span":{"begin":235,"end":240},"obj":"NCBITaxon:9606"},{"id":"T2624","span":{"begin":281,"end":286},"obj":"SO_EXT:0000704"},{"id":"T2625","span":{"begin":287,"end":293},"obj":"GO_EXT:biological_growth_entity_or_process"},{"id":"T2626","span":{"begin":364,"end":387},"obj":"SO_EXT:0000771"},{"id":"T2627","span":{"begin":389,"end":392},"obj":"SO_EXT:0000771"},{"id":"T2628","span":{"begin":413,"end":416},"obj":"SO_EXT:0000771"},{"id":"T2629","span":{"begin":417,"end":424},"obj":"SO_EXT:0001023"},{"id":"T2630","span":{"begin":458,"end":461},"obj":"SO_EXT:0000771"},{"id":"T2631","span":{"begin":505,"end":510},"obj":"NCBITaxon:10088"},{"id":"T2632","span":{"begin":511,"end":517},"obj":"GO_EXT:biological_growth_entity_or_process"},{"id":"T2633","span":{"begin":530,"end":533},"obj":"SO_EXT:0000771"},{"id":"T2634","span":{"begin":653,"end":660},"obj":"SO_EXT:0000704"},{"id":"T2635","span":{"begin":661,"end":669},"obj":"SO_EXT:sequence_variant_entity"},{"id":"T2636","span":{"begin":701,"end":704},"obj":"SO_EXT:0000771"},{"id":"T2637","span":{"begin":770,"end":775},"obj":"SO_EXT:0000704"},{"id":"T2638","span":{"begin":813,"end":818},"obj":"SO_EXT:0000704"},{"id":"T2639","span":{"begin":868,"end":875},"obj":"SO_EXT:0000704"},{"id":"T2640","span":{"begin":876,"end":885},"obj":"SO_EXT:sequence_variation_entity_or_quality"},{"id":"T2641","span":{"begin":1012,"end":1023},"obj":"SO_EXT:0000704"},{"id":"T2642","span":{"begin":1034,"end":1038},"obj":"NCBITaxon:10088"},{"id":"T2643","span":{"begin":1160,"end":1163},"obj":"SO_EXT:0000771"},{"id":"T2644","span":{"begin":1232,"end":1235},"obj":"SO_EXT:0000771"},{"id":"T2645","span":{"begin":1236,"end":1243},"obj":"SO_EXT:0001023"},{"id":"T2646","span":{"begin":1297,"end":1312},"obj":"SO_EXT:0001645"},{"id":"T2647","span":{"begin":1305,"end":1312},"obj":"CHEBI_SO_EXT:molecular_indicator_or_label_or_marker_or_tag"},{"id":"T2648","span":{"begin":1321,"end":1328},"obj":"SO_EXT:0000704"},{"id":"T2649","span":{"begin":1670,"end":1676},"obj":"CHEBI_SO_EXT:molecular_indicator_or_label_or_marker_or_tag"},{"id":"T2650","span":{"begin":1708,"end":1712},"obj":"PATO_UBERON_EXT:male_or_bearer_of_maleness"},{"id":"T2651","span":{"begin":1713,"end":1717},"obj":"NCBITaxon:10088"},{"id":"T2652","span":{"begin":1741,"end":1748},"obj":"SO_EXT:0001023"},{"id":"T2653","span":{"begin":1891,"end":1896},"obj":"NCBITaxon:10088"},{"id":"T2654","span":{"begin":1897,"end":1900},"obj":"SO_EXT:0000771"},{"id":"T2655","span":{"begin":1939,"end":1945},"obj":"GO_EXT:biological_growth_entity_or_process"},{"id":"T2656","span":{"begin":1990,"end":1995},"obj":"NCBITaxon:10088"},{"id":"T2657","span":{"begin":2023,"end":2029},"obj":"GO_EXT:biological_growth_entity_or_process"},{"id":"T2658","span":{"begin":2059,"end":2067},"obj":"SO_EXT:sequence_deletion_entity_or_process"},{"id":"T2659","span":{"begin":2095,"end":2105},"obj":"GO:0010467"},{"id":"T2660","span":{"begin":2113,"end":2118},"obj":"PR_EXT:000015393"},{"id":"T2661","span":{"begin":2120,"end":2154},"obj":"PR_EXT:000015393"},{"id":"T2662","span":{"begin":2134,"end":2142},"obj":"GO_EXT:0005125"},{"id":"T2663","span":{"begin":2134,"end":2152},"obj":"GO:0019221"},{"id":"T2664","span":{"begin":2156,"end":2160},"obj":"SO_EXT:0000704"},{"id":"T2665","span":{"begin":2182,"end":2187},"obj":"PR_EXT:000015393"},{"id":"T2666","span":{"begin":2253,"end":2258},"obj":"PR_EXT:000015393"},{"id":"T2667","span":{"begin":2258,"end":2259},"obj":"SO_EXT:sequence_nullness_or_absence"},{"id":"T2668","span":{"begin":2260,"end":2261},"obj":"SO_EXT:sequence_nullness_or_absence"},{"id":"T2669","span":{"begin":2271,"end":2276},"obj":"NCBITaxon:10088"},{"id":"T2670","span":{"begin":2358,"end":2361},"obj":"SO_EXT:0000771"},{"id":"T2671","span":{"begin":2442,"end":2445},"obj":"SO_EXT:0000771"},{"id":"T2672","span":{"begin":2461,"end":2464},"obj":"SO_EXT:0000771"},{"id":"T2673","span":{"begin":2527,"end":2535},"obj":"SO_EXT:sequence_alteration_process"},{"id":"T2674","span":{"begin":2563,"end":2567},"obj":"SO_EXT:0000704"},{"id":"T2675","span":{"begin":2616,"end":2624},"obj":"SO_EXT:sequence_alteration_entity_or_process"},{"id":"T2676","span":{"begin":2752,"end":2756},"obj":"SO_EXT:0000704"},{"id":"T2677","span":{"begin":2799,"end":2810},"obj":"CHEBI_EXT:biochemical"},{"id":"T2678","span":{"begin":2814,"end":2823},"obj":"GO:0008152"},{"id":"T2679","span":{"begin":2843,"end":2850},"obj":"SO_EXT:0000704"},{"id":"T2680","span":{"begin":2893,"end":2899},"obj":"GO_EXT:biological_growth_entity_or_process"},{"id":"T2681","span":{"begin":2963,"end":2968},"obj":"PR_EXT:000015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use of mouse models has provided valuable insight into the etiology of monogenic syndromes caused by single gene mutations. However, such models do not mimic the genetic complexity of disease traits commonly seen in the human population. Complex traits, such as polygenic growth and obesity are influenced by the small to moderate direct effects of quantitative trait loci (QTL), epistasis between QTL alleles, environmental perturbations and QTL-environment interactions.\nTo date numerous mouse growth and obesity QTL have been localized [1,2], however, little progress has been made in determining the specific nature of the underlying genetic variants. One resource used to fine map QTL are congenic strains which are designed to convert a complex polygenic trait into one that is mono- or oligogenic. This is accomplished by eliminating segregating genetic variation outside the locus of interest and reducing the environmental variation influencing a trait by characterizing large numbers of genetically identical mice. The Complex Trait Consortium (CTC) considers congenic analysis an excellent method to confirm and subsequently fine map QTL [3].\nTraditionally, congenic strains are developed by introgressing QTL alleles from a donor strain, whose boundaries are defined by genetic markers, on the genetic background of a recipient strain via 10 backcrosses [4]. While technically straightforward this is a time consuming endeavor taking over three years to construct a single strain. The speed congenic approach is an alternative to this lengthy process and can reduce the number of required backcrosses from 10 to five [5,6]. This strategy uses marker-assisted selection to identify male mice inheriting fewer donor alleles, than expected on average, during each backcross. Numerous traditionally developed and speed congenics have been used to successfully isolate mouse QTL for a wide array of traits, including growth and obesity [7-13].\nThe C57Bl/6J-hg/hg (HG) mouse is a model of systemic overgrowth resulting from a spontaneous deletion on MMU10, which eliminates expression of the Socs2 (suppressor of cytokine signaling 2) gene [14-16]. The role of Socs2 in the HG phenotype was confirmed by an independently engineered Socs2-/- knockout mouse which shared a number of phenotypes in common with HG, including gigantism [17].\nQTL which alter the phenotypic effects of another locus are referred to as modifier QTL [18]. Modifier QTL have been mapped for numerous traits and in these studies the modified locus is typically a known gene containing a spontaneously arisen or engineered mutation with major phenotypic effects, such as hg [11,19-21]. Epistasis forms the basis of these interactions, implying that the known gene and its modifiers are members of the same biochemical or metabolic pathway. Different genetic backgrounds have been shown to modify the growth-enhancing effects of hg [15,22]. Since the primary function of Socs2 is to negatively regulate growth hormone (Gh) [23], it is likely these background effects are the result of polymorphism influencing interactions between members of the Gh signaling pathway. Thus, identification of hg modifiers has the potential to uncover novel members of metabolically important pathways or previously unknown convergences between pathways.\nAs an initial step, growth and carcass composition QTL were identified in a cross between CAST/EiJ (CAST) and HG [24]. In the F2, mice homozygous for the hg deletion (hg/hg) and wild type (+/+) were assayed and hg modifier QTL were defined as those either absent in +/+ mice and segregating in hg/hg or loci with altered gene action dependent on background. Four hg modifier QTL (Wg2 on MMU2, Carfhg2 on MMU9, Carp2 on MMU11 and Feml3 on MMU17) were identified, along with 12 additional QTL (Q1Ucd1 on MMU1 (which only reached a suggestive level of statistical significance), Wg1, Carp1, Cara1 and Feml1 on MMU2, Carfhg1 on MMU5, Wg3 on MMU8, Feml2 on MMU9, Wg4 and Cara2 on MMU11 and Carp3 and Cara3 on MMU17) whose expression was independent of the hg locus.\nIn the current study we have developed speed congenic strains to isolate all the aforementioned QTL. Single strains on an HG background were created for each chromosomal region outside of MMU2, while a comprehensive panel of overlapping strains with identical donor regions on both B6 and HG backgrounds were developed for MMU2. The MMU2 panel was developed to confirm the presence of multiple QTL and test for QTL-hg interactions. Additionally, based on the knowledge of potential QTL-hg interactions, genes involved in Gh signaling, whose genomic location overlapped hg modifier QTL on MMU2, 9, 11 and 17, were sequenced."}

{"project":"craft-ca-core-dev","denotations":[{"id":"T2553","span":{"begin":22,"end":27},"obj":"NCBITaxon:10088"},{"id":"T2554","span":{"begin":90,"end":95},"obj":"SO:0000704"},{"id":"T2555","span":{"begin":123,"end":127},"obj":"SO:0000704"},{"id":"T2556","span":{"begin":177,"end":184},"obj":"SO:0000704"},{"id":"T2557","span":{"begin":235,"end":240},"obj":"NCBITaxon:9606"},{"id":"T2558","span":{"begin":281,"end":286},"obj":"SO:0000704"},{"id":"T2559","span":{"begin":364,"end":387},"obj":"SO:0000771"},{"id":"T2560","span":{"begin":389,"end":392},"obj":"SO:0000771"},{"id":"T2561","span":{"begin":413,"end":416},"obj":"SO:0000771"},{"id":"T2562","span":{"begin":417,"end":424},"obj":"SO:0001023"},{"id":"T2563","span":{"begin":458,"end":461},"obj":"SO:0000771"},{"id":"T2564","span":{"begin":505,"end":510},"obj":"NCBITaxon:10088"},{"id":"T2565","span":{"begin":530,"end":533},"obj":"SO:0000771"},{"id":"T2566","span":{"begin":653,"end":660},"obj":"SO:0000704"},{"id":"T2567","span":{"begin":701,"end":704},"obj":"SO:0000771"},{"id":"T2568","span":{"begin":770,"end":775},"obj":"SO:0000704"},{"id":"T2569","span":{"begin":813,"end":818},"obj":"SO:0000704"},{"id":"T2570","span":{"begin":868,"end":875},"obj":"SO:0000704"},{"id":"T2571","span":{"begin":1012,"end":1023},"obj":"SO:0000704"},{"id":"T2572","span":{"begin":1034,"end":1038},"obj":"NCBITaxon:10088"},{"id":"T2573","span":{"begin":1160,"end":1163},"obj":"SO:0000771"},{"id":"T2574","span":{"begin":1232,"end":1235},"obj":"SO:0000771"},{"id":"T2575","span":{"begin":1236,"end":1243},"obj":"SO:0001023"},{"id":"T2576","span":{"begin":1297,"end":1312},"obj":"SO:0001645"},{"id":"T2577","span":{"begin":1321,"end":1328},"obj":"SO:0000704"},{"id":"T2578","span":{"begin":1713,"end":1717},"obj":"NCBITaxon:10088"},{"id":"T2579","span":{"begin":1741,"end":1748},"obj":"SO:0001023"},{"id":"T2580","span":{"begin":1891,"end":1896},"obj":"NCBITaxon:10088"},{"id":"T2581","span":{"begin":1897,"end":1900},"obj":"SO:0000771"},{"id":"T2582","span":{"begin":1990,"end":1995},"obj":"NCBITaxon:10088"},{"id":"T2583","span":{"begin":2095,"end":2105},"obj":"GO:0010467"},{"id":"T2584","span":{"begin":2113,"end":2118},"obj":"PR:000015393"},{"id":"T2585","span":{"begin":2120,"end":2154},"obj":"PR:000015393"},{"id":"T2586","span":{"begin":2134,"end":2152},"obj":"GO:0019221"},{"id":"T2587","span":{"begin":2156,"end":2160},"obj":"SO:0000704"},{"id":"T2588","span":{"begin":2182,"end":2187},"obj":"PR:000015393"},{"id":"T2589","span":{"begin":2253,"end":2258},"obj":"PR:000015393"},{"id":"T2590","span":{"begin":2271,"end":2276},"obj":"NCBITaxon:10088"},{"id":"T2591","span":{"begin":2358,"end":2361},"obj":"SO:0000771"},{"id":"T2592","span":{"begin":2442,"end":2445},"obj":"SO:0000771"},{"id":"T2593","span":{"begin":2461,"end":2464},"obj":"SO:0000771"},{"id":"T2594","span":{"begin":2563,"end":2567},"obj":"SO:0000704"},{"id":"T2595","span":{"begin":2752,"end":2756},"obj":"SO:0000704"},{"id":"T2596","span":{"begin":2814,"end":2823},"obj":"GO:0008152"},{"id":"T2597","span":{"begin":2843,"end":2850},"obj":"SO:0000704"},{"id":"T2598","span":{"begin":2963,"end":2968},"obj":"PR:000015393"},{"id":"T2599","span":{"begin":3138,"end":3158},"obj":"GO:0060396"},{"id":"T2600","span":{"begin":3243,"end":3256},"obj":"GO:0008152"},{"id":"T2601","span":{"begin":3360,"end":3367},"obj":"UBERON:0008979"},{"id":"T2602","span":{"begin":3380,"end":3383},"obj":"SO:0000771"},{"id":"T2603","span":{"begin":3459,"end":3463},"obj":"NCBITaxon:10088"},{"id":"T2604","span":{"begin":3552,"end":3555},"obj":"SO:0000771"},{"id":"T2605","span":{"begin":3599,"end":3603},"obj":"NCBITaxon:10088"},{"id":"T2606","span":{"begin":3650,"end":3654},"obj":"SO:0000704"},{"id":"T2607","span":{"begin":3704,"end":3707},"obj":"SO:0000771"},{"id":"T2608","span":{"begin":3816,"end":3819},"obj":"SO:0000771"},{"id":"T2609","span":{"begin":4046,"end":4056},"obj":"GO:0010467"},{"id":"T2610","span":{"begin":4186,"end":4189},"obj":"SO:0000771"},{"id":"T2611","span":{"begin":4484,"end":4487},"obj":"SO:0000771"},{"id":"T2612","span":{"begin":4501,"end":4504},"obj":"SO:0000771"},{"id":"T2613","span":{"begin":4572,"end":4575},"obj":"SO:0000771"},{"id":"T2614","span":{"begin":4593,"end":4598},"obj":"SO:0000704"},{"id":"T2615","span":{"begin":4611,"end":4623},"obj":"GO:0060396"},{"id":"T2616","span":{"begin":4631,"end":4638},"obj":"SO:0001026"},{"id":"T2617","span":{"begin":4671,"end":4674},"obj":"SO:0000771"}],"text":"Background\nThe use of mouse models has provided valuable insight into the etiology of monogenic syndromes caused by single gene mutations. However, such models do not mimic the genetic complexity of disease traits commonly seen in the human population. Complex traits, such as polygenic growth and obesity are influenced by the small to moderate direct effects of quantitative trait loci (QTL), epistasis between QTL alleles, environmental perturbations and QTL-environment interactions.\nTo date numerous mouse growth and obesity QTL have been localized [1,2], however, little progress has been made in determining the specific nature of the underlying genetic variants. One resource used to fine map QTL are congenic strains which are designed to convert a complex polygenic trait into one that is mono- or oligogenic. This is accomplished by eliminating segregating genetic variation outside the locus of interest and reducing the environmental variation influencing a trait by characterizing large numbers of genetically identical mice. The Complex Trait Consortium (CTC) considers congenic analysis an excellent method to confirm and subsequently fine map QTL [3].\nTraditionally, congenic strains are developed by introgressing QTL alleles from a donor strain, whose boundaries are defined by genetic markers, on the genetic background of a recipient strain via 10 backcrosses [4]. While technically straightforward this is a time consuming endeavor taking over three years to construct a single strain. The speed congenic approach is an alternative to this lengthy process and can reduce the number of required backcrosses from 10 to five [5,6]. This strategy uses marker-assisted selection to identify male mice inheriting fewer donor alleles, than expected on average, during each backcross. Numerous traditionally developed and speed congenics have been used to successfully isolate mouse QTL for a wide array of traits, including growth and obesity [7-13].\nThe C57Bl/6J-hg/hg (HG) mouse is a model of systemic overgrowth resulting from a spontaneous deletion on MMU10, which eliminates expression of the Socs2 (suppressor of cytokine signaling 2) gene [14-16]. The role of Socs2 in the HG phenotype was confirmed by an independently engineered Socs2-/- knockout mouse which shared a number of phenotypes in common with HG, including gigantism [17].\nQTL which alter the phenotypic effects of another locus are referred to as modifier QTL [18]. Modifier QTL have been mapped for numerous traits and in these studies the modified locus is typically a known gene containing a spontaneously arisen or engineered mutation with major phenotypic effects, such as hg [11,19-21]. Epistasis forms the basis of these interactions, implying that the known gene and its modifiers are members of the same biochemical or metabolic pathway. Different genetic backgrounds have been shown to modify the growth-enhancing effects of hg [15,22]. Since the primary function of Socs2 is to negatively regulate growth hormone (Gh) [23], it is likely these background effects are the result of polymorphism influencing interactions between members of the Gh signaling pathway. Thus, identification of hg modifiers has the potential to uncover novel members of metabolically important pathways or previously unknown convergences between pathways.\nAs an initial step, growth and carcass composition QTL were identified in a cross between CAST/EiJ (CAST) and HG [24]. In the F2, mice homozygous for the hg deletion (hg/hg) and wild type (+/+) were assayed and hg modifier QTL were defined as those either absent in +/+ mice and segregating in hg/hg or loci with altered gene action dependent on background. Four hg modifier QTL (Wg2 on MMU2, Carfhg2 on MMU9, Carp2 on MMU11 and Feml3 on MMU17) were identified, along with 12 additional QTL (Q1Ucd1 on MMU1 (which only reached a suggestive level of statistical significance), Wg1, Carp1, Cara1 and Feml1 on MMU2, Carfhg1 on MMU5, Wg3 on MMU8, Feml2 on MMU9, Wg4 and Cara2 on MMU11 and Carp3 and Cara3 on MMU17) whose expression was independent of the hg locus.\nIn the current study we have developed speed congenic strains to isolate all the aforementioned QTL. Single strains on an HG background were created for each chromosomal region outside of MMU2, while a comprehensive panel of overlapping strains with identical donor regions on both B6 and HG backgrounds were developed for MMU2. The MMU2 panel was developed to confirm the presence of multiple QTL and test for QTL-hg interactions. Additionally, based on the knowledge of potential QTL-hg interactions, genes involved in Gh signaling, whose genomic location overlapped hg modifier QTL on MMU2, 9, 11 and 17, were sequenced."}