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Angelo Antonini Thaís Armangué Caroline Arquizan Scott Ayton William Banks Brenda Banwell Giuseppe Barisano Frederik Barkhof Karen Barlow Erin Beck Heleen Beckerman Alim Louis Benabid Selim Benbadis James Bernat Geert Jan Biessels Laurent Billot Niels Birbaumer Gretchen Birbeck Bradley Boeve Hayrunnisa Bolay Carsten Bonnemann Marie-Germaine Bousser Carol Brayne Xandra Owen Breakefield Alexis Brice Vera Bril Fabienne Brilot Matthijs Brouwer Adrian Budhram Jean Marc Burgunder Paolo Calabresi Bruce Campbell Cheryl Carcel Francisco Cardoso Sandra Morais Cardoso María Carmona-Iragui Jean-Laurent Casanova Neil Cashman Valeria Caso Fernando Cendes Piu Chan Andreas Charidimou Jeremy Chataway K Ray Chaudhuri Lei Chen Adriano Chiò Tanuja Chitnis Soo-jin Cho Shu-Ling Chong Leonid Churilov Paola Cinque Hans Clusmann Jeffrey Cohen Johnathan Cooper-Knock John Corboy Philippe Corcia Dennis Cordato Mario Cornejo-Olivas Irene Cortese Ana Sofia Costa Shelagh Coutts Maria L Cuadrado Merit Cudkowicz Russell C Dale Kristen Dams-O’Connor Yves Dauvilliers Mamede de Carvalho Bart De Strooper Marianne de Visser Joris de Wit Sophie Demeret Christel Depienne Pascal Derkinderen Jean-Francois Desaphy Orrin Devinsky David Devos Massimiliano Di Filippo Vincenzo Di Lazzaro Ramon Diaz-Arrastia Sulayman D Dib-hajj Hans-Christoph Diener Birger Victor Dieriks Ding Ding Ruth Dobson Matthew Dodd Qiang Dong Anne Ducros Jaroslaw Dulski John Duncan Alexandra Durr H Henrik Ehrsson David Eidelberg Alberto Espay Paolo Eusebi Amelia Evoli Margherita Fabbri Dongsheng Fan Forough Farrokhyar Alfonso Fasano Camille Fauchon Michael Fehlings Anthony Feinstein Eva Feldman Wuwei Feng Luigi Ferini Strambi Pierre-Olivier Fernagut Clarissa Ferrari Thalia Field Anthony Figaji Massimo Filippi Richard Finkel Urs Fischer Kathryn C Fitzgerald Agnès Fleury Jennifer Fogarty Brent Fogel Gary A Ford Juan Fortea Nick Fox Jacqueline French Giovanni Frisoni Kaiming G Fu Qiushi Fu Cynthia Gagnon Marialuisa Gandolfi Ziv Gan-Or Justo Garcĺa de Yébenes Angela Genge Marco Geraci Markus Glatzel Joseph Gleeson Jennifer Goldman Philip B Gorelick Rebecca Gottesman Mayank Goyal Francesc Graus Ribas Lea Grinberg James C Grotta James Guest Tanya Gurevich Mark Hallett Roy H Hamilton Adam Handel Graeme Hankey Oskar Hansson Orla Hardiman Michael Harhay David Hasan Nobutaka Hattori Jeffrey Hausdorff Yulei He Jacqueline S Hebert Michael Heckman Luke Henderson Bettina Henzi Amanda Heslegrave Michael Hill Marius Hoeper Günter Höglinger Reinhard Hohlfeld Rita Horvath Michael Howell Salvador Ibáñez Micó Sarosh Irani Muireann Irish Rigmor Højland Jensen Yong Ji Jiyao Jiang Liqun Jiao Peng Jin Anne Joutel Steven Julious Larry Junck Angela Kaindl Lorraine Kalia Anita Kamondi Takashi Kanda Kejal Kantarci Mark Kelson Zoe Kelson Ashvini Keshavan Pooja Khatri Joep Killestein Han-Joon Kim Helen Kim Jun-ichi Kira Christine Klein Benzi Kluger Martin Kocher Masatoshi Koga Daniel Kondziella Igor Koralnik Diana Krause Florian Krismer Kishore Kumar Shenghan Kuo Gert Kwakkel Jean-Charles Lambert Gert Jan Lammers Susan Landau Anthony Lang Simona Lattanzi Weidong Le Victoria Leavitt Christine Lebrun Frenay Edward Lee Min Ae Lee-Kirsch Andrew J Lees Robin Lemmens Elizabeth Leroux Johannes Levin Simon John Geoffrey Lewis David Liebeskind Shen-Yang Lim Hester Lingsma Ming Liu Tobias Loddenkemper Andres Lozano Madia Lozupone Liming Lu Wei Luo Andrew Maas Antoinette MaassenVanDenBrink R Loch Macdonald Pedro Machado Priyanka Madaan Walter Maetzler Melinda Magyari Shumei Man Mohammad Ali Mansournia Niklas Marklund Raúl Martínez-Fernández Sheila Martins Mario Masellis Clio Mavragani Mikael Mazighi Kimford Meador Martin I Meltzer Benedict Michael Timothy Miller Wendy Mitchell Biswadev Mitra John Mitrofanis Dimos-Dimitrios Mitsikostas Javadpour Mohsen Brit Mollenhauer Rikke Møller Xavier Montalban Teshamae Monteith Lewis Morgenstern Takeshi Morimoto Elena Moro Meg Morris Robert W Motl Tahseen Mozaffar Susanne Muehlschlegel Keith Muir Thomas Müller Praveen V Mummaneni Srikanth Muppidi Merrilee Needham Virginia Newcombe Thanh Nguyen Agneta Nordberg John Norrie Jose Obeso John O’Brien Neil Edward O’Connell Hirokazu Oguni Adesola Ogunniyi Michael Okun Verónica Olavarría Osamu Onodera Simona Orcesi Johanna Ospel Michael O’Sullivan Mayowa Owolabi Jacqueline Palace Lina Palaiodimou Elizabeth Emma Palmer Jeyaraj Pandian Deborah Pareto Lucilla Parnetti Mark Parsons Patrizio Pasqualetti Archana Patel Jean-Francois Payen Lillian Pazvakawambwa Ekaterina Pechenkova Yann Péréon Alexandra Pérez Emilio Perucca Piero Perucca Susanne Petri Dzung Pham Fredrik Piehl Yolande Pijnenburg Jennie Ponsford Lawrence Poree Kathleen Poston Marie-Claude Potier Andrea Domenico Pratico Christopher Price Federica Provini Alejandro Rabinstein Bianca Raffaelli Michael Rafii Yusuf Rajabally Tarek Rajji Sudarshini Ramanathan Olivier Rascol Sabrina Ravaglia Paola Rebora Irena Rektorová Uwe Reuter Katy Rezvani Marc Ribo Alan S Rigby Peter Ringleb Gabriël Rinkel Nilo Riva Chiara Robba Maria Rocca María Rodríguez-Oroz Michele Romoli Jonathan Rosand Clark Rosen Owen Ross Xavier Rossello Jeffrey Rothstein Peter Rothwell Àlex Rovira Christopher Rowe Annemieke J M Rozemuller Peter Rudge Ruediger Rupp Altaf Saadi Simona Sacco Mohammad Ali Sahraian Christian Saleh Cristina Sampaio Rujipat Samransamruajkit Ley Sander Else Charlotte Sandset Patrick Santens Naveed Sattar Ingrid E Scheffer Nicholas Schiff Julie Schneider Lon Schneider Christiane Schneider-Gold Andreas Schulze-Bonhage Giorgio Scivoletto Barry Seemungal James Sejvar Gustavo Sevlever Pankaj Sharma Christopher Shaw Pamela Shaw Kevin Sheth Fu-Dong Shi Holly A Shill Ashkan Shoamanesh Andew Siderowf Vincenzo Silani Robert Silbergleit Andrew Singleton Pyry Sipilä Claudia Sommer Lili Song Maria Grazia Spillantini Tara L Spires-Jones Luciano A Sposato Werner Stenzel Katharina Stibrant Sunnerhagen A Jon Stoessl Pasquale Striano Stephen Strittmatter José Ignacio Suárez Thyagarajan Subramanian John Suckling Antonio Suppa Heidi Sutherland Shigeaki Suzuki Kevin Talbot Eng-King Tan Keiko Tanaka Cristina Tassorelli Hudson Gerry Taylor John-Paul Taylor Hélio Afonso Teive Charlotte Teunissen Avner Thaler Madhav Thambisetty Sudhin Thayyil Eric Peter Thelin Roland Thijs Götz Thomalla Alan Thompson Lars Timmermann Mathias Toft Antonio Toscano Kazunori Toyoda Anthony Traboulsee Georgios Tsivgoulis Shoji Tsuji Arianna Tucci Guillaume Turc Andrew Udy Osamu Uemura Akiyuki Uzawa Pedro Valdes-Sosa Leonard van den Berg Ingrid A F van der Mei Anneke Van der Walt Ruben van Eijk Thomas Van Essen Sampsa Vanhatalo Lidia Vaqué Rosario Vasta Vinata Vedam-Mai Elisa Vegezzi Marie Vidailhet Aleksandar Videnovic Sylvia Villeneuve John Vissing Antonio Vitobello Sabine Voigt Victor Volovici Ostoja (Steve) Vucic Waqar Waheed Mitch Wallin Sebastian Walsh Jithangi Wanigasinghe Steven Warach Joanna Wardlaw Jason Warren Mike Wattjes Rimona Weil Paul Weiss Patrick Wen Marieke Wermer A Clinton White Robyn N Whitney Heinz S Wiendl Tissa Wijeratne Edward Wild Zachary Wills Nina Wilson Andrea Winkler Thomas Wisniewski Max Wiznitzer Nicole Wolf Jean Woo Anna Woodbury John A Woolmore Selina Wray Weili Xu Shadi Yaghi Pinar Yalinay-Dikmen Chuanzhu Yan Riqiang Yan Pengfei Yang Yin Yang Gerald W Zamponi Phyllis Zee Inga Zerr Xin Zhang Zhentao Zhang Dong Zhou Wendy Ziai

The food enzyme rennet paste containing chymosin (EC 3.4.23.4), pepsin A (EC 3.4.23.1) and triacylglycerol lipase (triacylglycerol acylhydrolase, EC 3.1.1.3) is prepared from the abomasum of suckling goats, lambs and calves by Caglificio Clerici S.p.A. The food enzyme is intended to be used in milk processing for cheese production. As no concerns arise from the animal source of the food enzyme, from its manufacture, and based on the history of safe use and consumption, the Panel considers that toxicological data were not required and no exposure assessment was necessary. On the basis of literature data, the Panel considers that, under the intended conditions of use, the risk of allergic sensitisation and elicitation reactions by dietary exposure could not be excluded, but the likelihood is considered to be low. Based on the data provided, the Panel concludes that this food enzyme does not give rise to safety concerns under the intended conditions of use.

Abstract This study was conducted to compare the effects of adding sodium butyrate (SB), medium-chain fatty acids (MCFAs), or n-3 polyunsaturated fatty acids (n-3 PUFAs) to the diet of sows during late gestation and lactation on the reproductive performance of sows and the growth performance and intestinal health of suckling piglets. Twenty-four sows (Landrace × Large-White hybrid; third parity; 200 ± 15 kg) were randomly assigned to receive 1 of 4 diets: basal diet (control group), basal diet + 1 g SB/kg (SB group), basal diet + 7.75 g MCFA/kg (MCFA group), or basal diet + 68.2 g n-3 PUFA/kg (n-3 PUFA group). The experiment began on day 85 of gestation and ended day 22 of lactation. Colostrum samples were collected from each sow. After the experiment, blood and tissue samples were collected from 1 randomly selected piglet. The results showed that the weaning-to-estrus interval of sows in the SB, MCFA, and n-3 PUFA groups was shorter than that of sows in the control group (P < 0.05). The incidence of diarrhea in suckling piglets in the SB, MCFA, and n-3 PUFA groups was lower than that of piglets in the control group (P < 0.05). The fat, protein, IgA, IgG, and IgM concentration in colostrum from sows increased following dietary supplementation with SB, MCFA, or n-3 PUFA (P < 0.05). Comparison with the control group, the mRNA expression of claudin-1, zona occludens 1, and interleukin-10 increased in the jejunum mucosa of suckling piglets in the SB, MCFA, and n-3 PUFA groups, while that of TLR4 decreased (P < 0.05). Compared with the control group, the Chao1 and ACE indexes of microbial flora in the colon contents of piglets in the SB, MCFA, and MCFA groups increased (P < 0.05), while the relative abundance of Firmicutes, Actinobacteria, and Synergistetes decreased at the phylum level (P < 0.05). In conclusion, during late pregnancy and lactation, dietary SB supplementation had a greater effect on intestinal health and caused a greater decrease in preweaning mortality of suckling piglets than did dietary MCFA or n-3 PUFA supplementation; dietary MCFA supplementation shortened the weaning-to-estrus interval of sows to a greater extent than did dietary SB or n-3 PUFA supplementation; and dietary n-3 PUFA supplementation increased the fat and protein content in the colostrum to the greatest extent.

Background: DNA methylation is an epigenetic mechanism central to development and maintenance of complex mammalian tissues, but our understanding of its role in intestinal development is limited. Results: We use whole genome bisulfite sequencing, and find that differentiation of mouse colonic intestinal stem cells to intestinal epithelium is not associated with major changes in DNA methylation. However, we detect extensive dynamic epigenetic changes in intestinal stem cells and their progeny during the suckling period, suggesting postnatal epigenetic development in this stem cell population. We find that postnatal DNA methylation increases at 3' CpG islands (CGIs) correlate with transcriptional activation of glycosylation genes responsible for intestinal maturation. To directly test whether 3' CGI methylation regulates transcription, we conditionally disrupted two major DNA methyltransferases, Dnmt1 or Dnmt3a, in fetal and adult intestine. Deficiency of Dnmt1 causes severe intestinal abnormalities in neonates and disrupts crypt homeostasis in adults, whereas Dnmt3a loss was compatible with intestinal development. These studies reveal that 3' CGI methylation is functionally involved in the regulation of transcriptional activation in vivo, and that Dnmt1 is a critical regulator of postnatal epigenetic changes in intestinal stem cells. Finally, we show that postnatal 3' CGI methylation and associated gene activation in intestinal epithelial cells are significantly altered by germ-free conditions. Conclusions: Our results demonstrate that the suckling period is critical for epigenetic development of intestinal stem cells, with potential important implications for lifelong gut health, and that the gut microbiome guides and/or facilitates these postnatal epigenetic processes.

Social behavior is a key component of pig welfare on farms, but little is known on the development of social behaviors in piglets. This study aimed to explore social behaviors and identify early social styles in suckling piglets. Social behaviors of 68 piglets from 12 litters were scored continuously for 8 h per day at 21 and 42 days of age, and were included in a Hierarchical Clustering on Principal Components analysis to identify clusters of pigs with similar social styles. Social nosing represented 78% of all social interactions given. Three social styles were identified: low-solicited inactive animals (inactive), active animals (active), and highly-solicited avoiders (avoiders). Belonging to a cluster was independent of age, but was influenced by sex, with females being more represented in the ‘inactive’ cluster, and males in the ‘active’ cluster, whereas both sexes were equally represented in the ‘avoider’ cluster. Stability of piglets’ allocation to specific clusters over age was high in the ‘inactive’ (59%) and ‘active’ (65%) clusters, but low in the ‘avoider’ cluster (7%). Haptoglobin and growth rate were higher in ‘active’ than ‘inactive’ pigs, and intermediate in ‘avoiders’. Our findings suggest the existence of transient social styles in piglets, likely reflective of sexual dimorphism or health status.

Colostrum-derived passive immunity is central to the health, performance and welfare of neonatal beef-suckler calves, and economics of beef-farming enterprises. Compared to dairy calves, mainly Holstein-Friesian, there is much less research carried out on passive immunity and associated factors in beef calves. Thus, this review aimed to summarise and interpret published information and highlight areas requiring further research. The transfer of immunoglobulin G1 (IgG1) from blood to mammary secretions is greater for beef × dairy cows compared to most beef breed types. Considerable between-animal variance is evident in first-milking colostrum yield and immunoglobulin concentration of beef-suckler cow breed types. First-milking colostrum immunoglobulin concentrations are similar for within-quarter fractions and for the front and rear quarters of the udder. First-milking colostrum yield is higher for beef × dairy cows than beef × beef and purebred beef breeds, and higher for multiparous than primiparous cows, but generally colostrum immunoglobulin concentration is relatively similar for each of the respective categories. Consequently, colostrum immunoglobulin mass (volume × concentration) production in beef cows seems to be primarily limited by colostrum volume. The effect of maternal nutrition during late gestation on colostrum yield is not well documented; however, most studies provide evidence that colostrum immunoglobulin concentration is not adversely affected by under-nutrition. Factors that impinge upon the duration between birth and first suckling, including dam parity, udder and teat anatomy and especially dystocia, negatively impact on calf passive immunity. Colostrum immunoglobulin mass ingested relative to birth weight post-parturition is the most important variable determining calf passive immunity. Research indicates that feeding the beef calf a colostrum volume equivalent to 5% of birth weight shortly after parturition, with subsequent suckling of the dam (or a second feed) 6 to 8 h later, ensures adequate passive immunity, equivalent to a well-managed suckling situation. Within beef-suckler cow genotypes, calf passive immunity is similar for many common beef breeds, but is generally higher for calves from beef × dairy cows. Compared to older cows, calves from younger cows, especially primiparous animals, have lower serum immunoglobulin concentrations. Most studies have shown no adverse impact of maternal dietary restriction on calf passive immunity. The prevalence of failure of passive transfer (FPT) in beef calves varies considerably across studies depending on the test used, and what cut-off value is assumed or how it is classified. The accuracy and precision of methodologies used to determine immunoglobulin concentrations is concerning; caution is required in interpreting laboratory results regarding defining colostrum ‘quality’ and calf passive immune ‘status’. Further research is warranted on colostrum-related factors limiting passive immunity of beef calves, and on the validation of laboratory test cut-off points for determining FPT, based on their relationships with key health and performance measures.

At birth, when immune responses are insufficient, there begins the development of the defence capability against pathogens. Leptin and adiponectin, adipokines that are present in breast milk, have been shown to play a role in the regulation of immune responses. We report here, for the first time, the influence of in vivo adipokine supplementation on the intestinal immune system in early life. Suckling Wistar rats were daily supplemented with leptin (0·7 μg/kg per d, n 36) or adiponectin (35 μg/kg per d, n 36) during the suckling period. The lymphocyte composition, proliferation and cytokine secretion from mesenteric lymph node lymphocytes (on days 14 and 21), as well as intestinal IgA and IgM concentration (day 21), were evaluated. At day 14, leptin supplementation significantly increased the TCRαβ + cell proportion in mesenteric lymph nodes, in particular owing to an increase in the TCRαβ + CD8+ cell population. Moreover, the leptin or adiponectin supplementation promoted the early development CD8+ cells, with adiponectin being the only adipokine capable of enhancing the lymphoproliferative ability at the end of the suckling period. Although leptin decreased intestinal IgA concentration, it had a trophic effect on the intestine in early life. Supplementation of both adipokines modulated the cytokine profile during (day 14) and at the end (day 21) of the suckling period. These results suggest that leptin and adiponectin during suckling play a role in the development of mucosal immunity in early life.

•A mouse model was used to evaluate equine rotavirus vaccines.•G3 and G14 vaccines were both effective against G3 rotaviruses.•G14 vaccine was effective against G14 rotaviruses.•At least a G14 rotavirus should be included in a vaccine strain. Equine group A rotaviruses (RVAs) cause diarrhea in suckling foals. The dominant RVAs circulating among horses worldwide, including Japan, are G3P[12] and/or G14P[12] genotypes. Inactivated vaccines containing a G3P[12] RVA are commercially available in some countries for prevention of diarrhea caused by equine RVAs. However, there is no reported evidence whether vaccines containing a G3P[12] RVA are effective against G14P[12] RVAs or whether using a G14P[12] RVA results in a more effective vaccine. This study used a suckling mouse model to evaluate the effectiveness of inactivated vaccines containing G3P[12] (G3 vaccine) or G14P[12] (G14 vaccine) RVAs against G3P[12] and G14P[12] RVAs. Female mice were inoculated twice with G3 or G14 vaccines, and were then mated. After parturition, suckling mice were challenged with one of either two G3P[12] RVAs, two G14P[12] RVAs, or one G13P[18] RVA. After virus inoculation, suckling mice were observed for diarrhea, and the incidence rates of diarrhea in the vaccinated groups were compared with those in the non-vaccinated groups. Following G3P[12] RVA challenge, suckling mice in the G3 and G14 vaccinated groups had significantly lower rates of diarrhea incidence than did those in the non-vaccinated group, and the rates in the G3 vaccinated group tended to be lower than in the G14 vaccinated group. Following G14P[12] RVA challenge, suckling mice in the G14 vaccinated group had significantly lower rates of diarrhea incidence than did those in the non-vaccinated and G3 vaccinated groups. The G3 and G14 vaccines did not reduce the rate when challenged with the G13P[18] RVA. The mouse model showed that the G3 and G14 vaccines were both effective against G3P[12] RVAs, and that the G14 vaccine was effective against G14P[12] RVAs. These results suggest that at least a G14 RVA strain should be included in as a vaccine strain.

Myocarditis is considered a fatal form of foot-and-mouth disease (FMD) in suckling calves. In the present study, a total of 17 calves under 4 months of age and suspected clinically for FMD were examined for clinical lesions, respiratory rate, heart rate, and heart rhythm. Lesion samples, saliva, nasal swabs, and whole blood were collected from suspected calves and subjected to Sandwich ELISA and reverse transcription multiplex polymerase chain reaction (RT-mPCR) for detection and serotyping of FMD virus (FMDV). The samples were found to be positive for FMDV serotype “O”. Myocarditis was suspected in 6 calves based on tachypnoea, tachycardia, and gallop rhythm. Serum aspartate aminotransferase (AST), creatinine kinase myocardial band (CK-MB) and lactate dehydrogenase (LDH), and cardiac troponins (cTnI) were measured. Mean serum AST, cTn-I and LDH were significantly higher (P < 0.001) in < 2 months old FMD-infected calves showing clinical signs suggestive of myocarditis (264.833 ± 4.16; 11.650 ± 0.34 and 1213.33 ± 29.06) than those without myocarditis (< 2 months old: 110.00 ± 0.00, 0.06 ± 0.00, 1050.00 ± 0.00; > 2 months < 4 months: 83.00 ± 3.00, 0.05 ± 0.02, 1159.00 ± 27.63) and healthy control groups (< 2 months old: 67.50 ± 3.10, 0.047 ± 0.01, 1120.00 ± 31.62; > 2 months < 4 months: 72.83 ± 2.09, 0.47 ± 0.00, 1160.00 ± 18.44). However, mean serum CK-MB did not differ significantly amongst the groups. Four calves under 2 months old died and a necropsy revealed the presence of a pathognomic gross lesion of the myocardial form of FMD known as “tigroid heart”. Histopathology confirmed myocarditis. This study also reports the relevance of clinical and histopathological findings and biochemical markers in diagnosing FMD-related myocarditis in suckling calves.

Meat intramuscular fat (IMF) contributes to meat quality and consumer acceptance. Molecular events that occur during IMF deposition and the identification of genes that are differentially expressed during this process are important to the design of an optimal nutrition plan for animals. In the present study, we examined the effect of the forage type (grazing vs. hay pasture) fed to ewes and the effect of lamb sex on the LM fatty acid (FA) profile and gene expression of suckling lambs (10 to 12 kg of BW at slaughter); ewes received pasture hay (PH) or grazed pasture (GRE). Forage type had a significant effect on IMF FA profile. Ewes grazing green forage (GRE) promoted the formation and deposition of vaccenic acid (C18:1n-7), CLA, and PUFA n-3 in LM from their suckling lambs (P < 0.05). We found that forage type affected the expression of the sterol regulatory element binding transcription factor 1 (SREBF1) gene in females. However, in males, it modulated stearoyl CoA desaturase (SCD) gene expression (P < 0.05). Moreover, our results showed that females, independent of the diet of the ewes (PH or GRE), are predisposed to develop fat and to upregulate the expression of key genes of transcriptional factors PPARA, CEBPB, SREBF1, and lipoprotein lipase (LPL) and SCD (P < 0.05). The data suggest that SREBF1, SCD, and most likely CEBPB gene expression in young suckling lambs is modulated by both lamb sex and forage type fed to ewes. Fatty acid indicators PUFA, n-6/n-3, CLA, and SFA are closely related to LPL, SCD, PPARA, and CEBPB gene expression depending on animal sex or the diet of ewes. This study suggests that grazing pasture affects FA composition promoting greater vaccenic, CLA, and total PUFA n-3 FA in female and male suckling lambs, and it is mediated through the regulation of lipogenic enzyme expression.