Obesity during Pregnancy in the Horse: Effect on Term Placental Structure and Gene Expression, as Well as Colostrum and Milk Fatty Acid Concentration.
Abstract: In horses, the prevalence of obesity is high and associated with serious metabolic pathologies. Being a broodmare has been identified as a risk factor for obesity. In other species, maternal obesity is known to affect the development of the offspring. This article is a follow-up study of previous work showing that Obese mares (O, n = 10, body condition score > 4.25 at insemination) were more insulin resistant and presented increased systemic inflammation during pregnancy compared to Normal mares (N, n = 14, body condition score < 4 at insemination). Foals born to O mares were more insulin-resistant, presented increased systemic inflammation, and were more affected by osteoarticular lesions. The objective of the present study was to investigate the effect of maternal obesity on placental structure and function, as well as the fatty acid profile in the plasma of mares and foals, colostrum, and milk until 90 days of lactation, which, to our knowledge, has been poorly studied in the horse. Mares from both groups were fed the same diet during pregnancy and lactation. During lactation, mares were housed in pasture. A strong heat wave, followed by a drought, occurred during their 2nd and 3rd months of lactation (summer of 2016 in the Limousin region, France). In the present article, term placental morphometry, structure (stereology), and gene expression (RT-qPCR, genes involved in nutrient transport, growth, and development, as well as vascularization) were studied. Plasma of mares and their foals, as well as colostrum and milk, were sampled at birth, 30 days, and 90 days of lactation. The fatty acid composition of these samples was measured using gas chromatography. No differences between the N and O groups were observed for term placental morphometry, structure, or gene expression. No difference in plasma fatty acid composition was observed between groups in mares. The plasma fatty acid profile of O foals was more pro-inflammatory and indicated an altered placental lipid metabolism between birth and 90 days of age. These results are in line with the increased systemic inflammation and altered glucose metabolism observed until 18 months of age in this group. The colostrum fatty acid profile of O mares was more pro-inflammatory and indicated an increased transfer and/or desaturation of long-chain fatty acids. Moreover, O foals received a colostrum poorer in medium-chain saturated fatty acid, a source of immediate energy for the newborn that can also play a role in immunity and gut microbiota development. Differences in milk fatty acid composition indicated a decreased ability to adapt to heat stress in O mares, which could have further affected the metabolic development of their foals. In conclusion, maternal obesity affected the fatty acid composition of milk, thus also influencing the foal's plasma fatty acid composition and likely participating in the developmental programming observed in growing foals.
Publication Date: 2023-12-04 PubMed ID: 38133242PubMed Central: PMC10748288DOI: 10.3390/vetsci10120691Google Scholar: Lookup
The Equine Research Bank provides access to a large database of publicly available scientific literature. Inclusion in the Research Bank does not imply endorsement of study methods or findings by Mad Barn.
- Journal Article
Summary
This research summary has been generated with artificial intelligence and may contain errors and omissions. Refer to the original study to confirm details provided. Submit correction.
This research article focuses on the impacts of maternal obesity in horses on placental structure, gene expression, and the fatty acid concentration in colostrum and milk. While the study found no noticeable changes to placental features, it observed an increased pro-inflammatory fatty acid profile in obese mares’ offspring, indicating an altered placental lipid metabolism. Evident was also a negative impact on colostrum and milk quality, which likely influenced the offspring’s metabolic development.
Understanding the Study
- The study followed up on previous conclusions indicating high insulin resistance and systemic inflammation in obese mares during pregnancy. By comparing these with normal mares, it was discovered that obesity might affect the health and growth of horse offspring.
- The research aims to dig deeper into the effects of maternal obesity on the structure and function of the placenta. It also seeks to explore the fatty acid profile in the plasma of mares and foals along with the colostrum and milk over 90 days of lactation – a component that had not been given much attention in horse studies.
- The pregnant mares in both the obese and normal groups were subjected to the same diet during pregnancy and lactation, and were kept in a pasture during lactation. The conditions during lactation included a heatwave and drought.
Methodology
- Researchers used various methods to examine term placental morphometry, structure, and gene expression – this included stereology and RT-qPCR, targeting genes involved in nutrient transport, growth, development, and vascularization.
- Numerous samples were collected during the study – plasma from mares and foals, and colostrum and milk – at birth, after 30 days, and after 90 days of lactation. The fatty acid composition of these samples was assessed using gas chromatography.
Key Findings
- No significant differences were noted in placental morphometry, structure, or gene expression between normal and obese mares.
- The plasma fatty acid profile of offspring from obese mares was more pro-inflammatory and indicated altered placental lipid metabolism up to the age of 90 days. This outcome is consistent with previous findings of increased systemic inflammation and altered glucose metabolism in this group until 18 months of age.
- The fatty acid profile of colostrum from obese mares was also more pro-inflammatory; it suggested a heightened transfer and/or desaturation of long-chain fatty acids.
- The milk from obese mares had a lesser quantity of medium-chain saturated fatty acid that serves as an immediate energy source for newborns and is critical in immunity and gut microbiota development.
- Obese mares also showed a lessened ability to adapt to heat stress, likely affecting the metabolic development of their foals.
Conclusion
- The study concludes that maternal obesity can influence the fatty acid composition in milk, which in turn might impact the plasma fatty acid composition in foals and has potential effects on their metabolic development.
Cite This Article
APA
Robles M, Rousseau-Ralliard D, Dubois C, Josse T, Nouveau É, Dahirel M, Wimel L, Couturier-Tarrade A, Chavatte-Palmer P.
(2023).
Obesity during Pregnancy in the Horse: Effect on Term Placental Structure and Gene Expression, as Well as Colostrum and Milk Fatty Acid Concentration.
Vet Sci, 10(12).
https://doi.org/10.3390/vetsci10120691 Publication
Researcher Affiliations
- BREED, Domaine de Vilvert, Université Paris Saclay, UVSQ, INRAE, 78350 Jouy en Josas, France.
- BREED, Ecole Nationale Vétérinaire d'Alfort, 94700 Maisons-Alfort, France.
- Institut Polytechnique Unilasalle, 76130 Mont-Saint-Aignan, France.
- BREED, Domaine de Vilvert, Université Paris Saclay, UVSQ, INRAE, 78350 Jouy en Josas, France.
- BREED, Ecole Nationale Vétérinaire d'Alfort, 94700 Maisons-Alfort, France.
- Institut Français du Cheval et de l'Equitation, Station Expérimentale de la Valade, 19370 Chamberet, France.
- BREED, Domaine de Vilvert, Université Paris Saclay, UVSQ, INRAE, 78350 Jouy en Josas, France.
- BREED, Ecole Nationale Vétérinaire d'Alfort, 94700 Maisons-Alfort, France.
- BREED, Domaine de Vilvert, Université Paris Saclay, UVSQ, INRAE, 78350 Jouy en Josas, France.
- BREED, Ecole Nationale Vétérinaire d'Alfort, 94700 Maisons-Alfort, France.
- BREED, Domaine de Vilvert, Université Paris Saclay, UVSQ, INRAE, 78350 Jouy en Josas, France.
- BREED, Ecole Nationale Vétérinaire d'Alfort, 94700 Maisons-Alfort, France.
- Institut Français du Cheval et de l'Equitation, Station Expérimentale de la Valade, 19370 Chamberet, France.
- BREED, Domaine de Vilvert, Université Paris Saclay, UVSQ, INRAE, 78350 Jouy en Josas, France.
- BREED, Ecole Nationale Vétérinaire d'Alfort, 94700 Maisons-Alfort, France.
- BREED, Domaine de Vilvert, Université Paris Saclay, UVSQ, INRAE, 78350 Jouy en Josas, France.
- BREED, Ecole Nationale Vétérinaire d'Alfort, 94700 Maisons-Alfort, France.
Grant Funding
- FOETALIM / Institut Français du Cheval et de l'Équitation
- FOETALIM / Fonds Eperon
Conflict of Interest Statement
The authors declare no conflict of interest.
References
This article includes 108 references
- Peugnet P, Robles M, Wimel L, Tarrade A, Chavatte-Palmer P. Management of the Pregnant Mare and Long-Term Consequences on the Offspring. Theriogenology 2016;86:99–109.
- Robles M, Hammer C, Staniar B, Chavatte-Palmer P. Nutrition of Broodmares. Vet. Clin. Equine Pract. 2021;37:177–205.
- Chavatte-Palmer P, Derisoud E, Robles M. Pregnancy and Placental Development in Horses: An Update. Domest. Anim. Endocrinol. 2022;79:106692.
- Derisoud E, Auclair-Ronzaud J, Palmer E, Robles M, Chavatte-Palmer P, Derisoud E, Auclair-Ronzaud J, Palmer E, Robles M, Chavatte-Palmer P. Female Age and Parity in Horses: How and Why Does It Matter?. Reprod. Fertil. Dev. 2021;34:52–116.
- Peugnet P, Wimel L, Duchamp G, Sandersen C, Camous S, Guillaume D, Dahirel M, Dubois C, Jouneau L, Reigner F. Enhanced or Reduced Fetal Growth Induced by Embryo Transfer into Smaller or Larger Breeds Alters Post-Natal Growth and Metabolism in Pre-Weaning Horses. PLoS ONE 2014;9:e102044.
- Peugnet P, Mendoza L, Wimel L, Duchamp G, Dubois C, Reigner F, Caudron I, Deliège B, Toquet M, Richard E. Longitudinal Study of Growth and Osteoarticular Status in Foals Born to Between-Breed Embryo Transfers. J. Equine Vet. Sci. 2016;37:24–38.
- Giussani D, Forhead A, Gardner D, Fletcher A, Allen W, Fowden A. Postnatal Cardiovascular Function after Manipulation of Fetal Growth by Embryo Transfer in the Horse. J. Physiol. 2003;547:67–76.
- Forhead A, Ousey J, Allen W, Fowden A. Postnatal Insulin Secretion and Sensitivity after Manipulation of Fetal Growth by Embryo Transfer in the Horse. J. Endocrinol. 2004;181:459–467.
- Tischner M, Klimczak M. The Development of Polish Ponies Born after Embryo Transfer to Large Recipients. Equine Vet. J. 1989;21:62–63.
- Kubiak J, Evans J, Potter G, Harms P, Jenkins W. Parturition in the Multiparous Mare Fed to Obesity. Equine Vet. Sci. 1988;8:135–140.
- Kubiak J, Evans J, Potter G, Harms P, Jenkins W. Milk Yield Composition in the Multiparous Mare Fed to Obesity. Equine Vet. Sci. 1991;11:158–162.
- Henneke D, Potter G, Kreider J. Body Condition during Pregnancy and Lactation and Reproductive Efficiency of Mares. Theriogenology 1984;21:897–909.
- Bradbery A.N, Coverdale J.A, Hammer C.J, Dunlap K.A, Leatherwood J.L, Satterfield M.C. Effect of Maternal Overnutrition on Predisposition to Insulin Resistance in the Foal: Foal Skeletal Muscle Development and Insulin Signaling. Domest. Anim. Endocrinol. 2021;77:106648.
- Wilsher S, Allen W. Effects of a Streptococcus Equi Infection—Mediated Nutritional Insult during Mid-Gestation in Primiparous Thoroughbred Fillies. Part 1: Placental and Fetal Development. Equine Vet. J. 2006;38:549–557.
- Peugnet P, Robles M, Mendoza L, Wimel L, Dubois C, Dahirel M, Guillaume D, Camous S, Berthelot V, Toquet M.-P. Effects of Moderate Amounts of Barley in Late Pregnancy on Growth, Glucose Metabolism and Osteoarticular Status of Pre-Weaning Horses. PLoS ONE 2015;10:e0122596.
- Robles M, Gautier C, Mendoza L, Peugnet P, Dubois C, Dahirel M, Lejeune J, Caudron I, Guenon I, Camous S. Maternal Nutrition during Pregnancy Affects Testicular and Bone Development, Glucose Metabolism and Response to Overnutrition in Weaned Horses up to Two Years. PLoS ONE 2017;12:e0169295.
- Robles M, Nouveau E, Gautier C, Mendoza L, Dubois C, Dahirel M, Lagofun B, Aubrière M, Lejeune J, Caudron I. Maternal Obesity Increases Insulin Resistance, Low-Grade Inflammation and Osteochondrosis Lesions in Foals and Yearlings until 18 Months of Age. PLoS ONE 2018;13:e0190309.
- Giles S, Rands S, Nicol C, Harris P. Obesity Prevalence and Associated Risk Factors in Outdoor Living Domestic Horses and Ponies. PeerJ 2014;2:e299.
- Potter S, Bamford N, Harris P, Bailey S. Prevalence of Obesity and Owners’ Perceptions of Body Condition in Pleasure Horses and Ponies in South-Eastern Australia. Aust. Vet. J. 2016;94:427–432.
- Hoffman C, Costa L, Freeman L. Survey of Feeding Practices, Supplement Use, and Knowledge of Equine Nutrition among a Subpopulation of Horse Owners in New England. J. Equine Vet. Sci. 2009;29:719–726.
- Furtado T, Perkins E, Pinchbeck G, McGowan C, Watkins F, Christley R. Exploring Horse Owners’ Understanding of Obese Body Condition and Weight Management in UK Leisure Horses. Equine Vet. J. 2021;53:752–762.
- Hoffman R.M, Boston R.C, Stefanovski D, Kronfeld D.S, Harris P.A. Obesity and Diet Affect Glucose Dynamics and Insulin Sensitivity in Thoroughbred Geldings. J. Anim. Sci. 2003;81:2333–2342.
- Siwinska N, Janus I, Zak-Bochenek A, Noszczyk-Nowak A. Influence of Obesity on Histological Tissue Structure of the Cardiovascular System in Horses. Animals 2022;12:732.
- Lovett A.L, Gilliam L.L, Sykes B.W, McFarlane D. Thromboelastography in Obese Horses with Insulin Dysregulation Compared to Healthy Controls. J. Vet. Intern. Med. 2022;36:1131–1138.
- Pearson W, Wood K, Stanley S, Macnicol J. Exploring Relationships between Body Condition Score, Body Fat, Activity Level and Inflammatory Biomarkers. J. Anim. Physiol. Anim. Nutr. 2018;102:1062–1068.
- Salinas C, Espinosa G, Morales N, Henríquez C, Morán G, Gajardo G, Burgos R.A, Uberti B. Evaluation of Peripheral Blood Polymorphonuclear Cell Functions after an Oral Carbohydrate Overload in Obese and Insulin Dysregulated Horses. Vet. Immunol. Immunopathol. 2022;250:110455.
- Frank N. Equine Metabolic Syndrome. Vet. Clin. North Am. Equine Pract. 2011;27:73–92.
- Alford P, Geller S, Richrdson B, Slater M, Honnas C, Foreman J, Robinson J, Messer M, Roberts M, Goble D. A Multicenter, Matched Case-Control Study of Risk Factors for Equine Laminitis. Prev. Vet. Med. 2001;49:209–222.
- Garber A, Hastie P, Murray J.-A. Factors Influencing Equine Gut Microbiota: Current Knowledge. J. Equine Vet. Sci. 2020;88:102943.
- Roth P, Stanley J, Chamoun-Emanuelli A, Whitfield-Cargile C, Coleman M. Fecal Extract from Obese Horses Induces an Inflammatory Response by Murine Macrophages in Vitro. Am. J. Vet. Res. 2022;83:419–425.
- Pratt-Phillips S, Munjizun A. Impacts of Adiposity on Exercise Performance in Horses. Anim. Open Access J. 2023;13:666.
- Jansson A, Gunnarsson V.Þ., Ringmark S, Ragnarsson S, Söderroos D, Ásgeirsson E, Jóhannsdóttir T.R, Liedberg C, Stefánsdóttir G.J. Increased Body Fat Content in Horses Alters Metabolic and Physiological Exercise Response, Decreases Performance, and Increases Locomotion Asymmetry. Physiol. Rep. 2021;9:e14824.
- Jaqueth A.L, Iwaniuk M.E, Burk A.O. Characterization of the Prevalence and Management of Over-Conditioned Ponies and Horses in Maryland. J. Equine Vet. Sci. 2018;68:26–32.
- Shrestha N, Ezechukwu H.C, Holland O.J, Hryciw D.H. Developmental Programming of Peripheral Diseases in Offspring Exposed to Maternal Obesity during Pregnancy. Am. J. Physiol.-Regul. Integr. Comp. Physiol. 2020;319:R507–R516.
- Kelly A.C, Powell T.L, Jansson T. Placental Function in Maternal Obesity. Clin. Sci. 2020;134:961–984.
- Schoonejans J.M, Ozanne S.E. Developmental Programming by Maternal Obesity: Lessons from Animal Models. Diabet. Med. 2021;38:e14694.
- Lecoutre S, Maqdasy S, Breton C. Maternal Obesity as a Risk Factor for Developing Diabetes in Offspring: An Epigenetic Point of View. World J. Diabetes 2021;12:366–382.
- Kislal S, Shook L.L, Edlow A.G. Perinatal Exposure to Maternal Obesity: Lasting Cardiometabolic Impact on Offspring. Prenat. Diagn. 2020;40:1109–1125.
- Rodgers A, Sferruzzi-Perri A.N. Developmental Programming of Offspring Adipose Tissue Biology and Obesity Risk. Int. J. Obes. 2021;45:1170–1192.
- Kong L, Chen X, Gissler M, Lavebratt C. Relationship of Prenatal Maternal Obesity and Diabetes to Offspring Neurodevelopmental and Psychiatric Disorders: A Narrative Review. Int. J. Obes. 2020;44:1981–2000.
- Menting M.D, van de Beek C, Mintjens S, Wever K.E, Korosi A, Ozanne S.E, Limpens J, Roseboom T.J, Hooijmans C, Painter R.C. The Link between Maternal Obesity and Offspring Neurobehavior: A Systematic Review of Animal Experiments. Neurosci. Biobehav. Rev. 2019;98:107–121.
- Derisoud E, Auclair-Ronzaud J, Rousseau-Ralliard D, Philau S, Aujean E, Durand A, Dahirel M, Charlier M, Boutinaud M, Wimel L. Maternal Age, Parity and Nursing Status at Fertilization Affects Postpartum Lactation Up to Weaning in Horses. J. Equine Vet. Sci. 2023;128:104868.
- Robles M, Loux S.C, de Mestre A.M, Chavatte-Palmer P. Environmental Constraints and Pathologies That Modulate Equine Placental Genes and Development. Reprod. Camb. Engl. 2022;163:R25–R38.
- Allen W, Wilsher S, Turnbull C, Stewart F, Ousey J, Rossdale P, Fowden A, Turnbull C, Stewart F, Ousey J. Influence of Maternal Size on Placental, Fetal and Postnatal Growth in the Horse. I. Development in Utero. Reproduction 2002;172:445–453.
- Robles M, Peugnet P, Dubois C, Piumi F, Jouneau L, Bouchez O, Aubrière M, Dahirel M, Aioun J, Wimel L. Placental Function and Structure at Term Is Altered in Broodmares Fed with Cereals from Mid-Gestation. Placenta 2018;64:44–52.
- Auclair-Ronzaud J, Jaffrézic F, Wimel L, Dubois C, Laloë D, Chavatte-Palmer P. Estimation of Milk Production in Suckling Mares and Factors Influencing Their Milk Yield. Animal 2022;16:100498.
- Reed M, Howard C, DE Yanés G. One-Stop Stereology: The Estimation of 3D Parameters Using Isotropic Rulers. J. Microsc. 2010;239:54–65.
- Chomczynski P, Sacchi N. Single-Step Method of RNA Isolation by Acid Guanidinium Thiocyanate-Phenol-Chloroform Extraction. Anal. Biochem. 1987;162:156–159.
- Tarrade A, Rousseau-Ralliard D, Aubrière M, Peynot N, Dahirel M, Bertrand-Michel J, Aguirre-Lavin T, Morel O, Beaujean N, Duranthon V. Sexual Dimorphism of the Feto-Placental Phenotype in Response to a High Fat and Control Maternal Diets in a Rabbit Model. PLoS ONE 2013;8:e83458.
- Folch J, Lees M, Stanley G.S. A Simple Method for the Isolation and Purification of Total Lipides from Animal Tissues. J. Biol. Chem. 1957;226:497–509.
- Morrison W.R, Smith L.M. Preparation of Fatty Acid Methyl Esters and Dimethylacetals from Lipids with Boron Fluoride–Methanol. J. Lipid Res. 1964;5:600–608.
- Rousseau D, Héliès-Toussaint C, Moreau D, Raederstorff D, Grynberg A. Dietary N-3 PUFAs Affect the Blood Pressure Rise and Cardiac Impairments in a Hyperinsulinemia Rat Model in Vivo. Am. J. Physiol.-Heart Circ. Physiol. 2003;285:H1294–H1302.
- Dubé E, Gravel A, Martin C, Desparois G, Moussa I, Ethier-Chiasson M, Forest J.-C, Giguère Y, Masse A, Lafond J. Modulation of Fatty Acid Transport and Metabolism by Maternal Obesity in the Human Full-Term Placenta1. Biol. Reprod. 2012;87:14.
- Tian L, Dong S.S, Hu J, Yao J.J, Yan P.S. The Effect of Maternal Obesity on Fatty Acid Transporter Expression and Lipid Metabolism in the Full-Term Placenta of Lean Breed Swine. J. Anim. Physiol. Anim. Nutr. 2018;102:e242–e253.
- Song L, Wang N, Peng Y, Sun B, Cui W. Placental Lipid Transport and Content in Response to Maternal Overweight and Gestational Diabetes Mellitus in Human Term Placenta. Nutr. Metab. Cardiovasc. Dis. 2022;32:692–702.
- Nogues P, Dos Santos E, Couturier-Tarrade A, Berveiller P, Arnould L, Lamy E, Grassin-Delyle S, Vialard F, Dieudonne M.-N. Maternal Obesity Influences Placental Nutrient Transport, Inflammatory Status, and Morphology in Human Term Placenta. J. Clin. Endocrinol. Metab. 2020;12:dgaa660.
- Rasool A, Mahmoud T, Mathyk B, Kaneko-Tarui T, Roncari D, White K.O, O’Tierney-Ginn P. Obesity Downregulates Lipid Metabolism Genes in First Trimester Placenta. Sci. Rep. 2022;12:19368.
- Napso T, Lean S.C, Lu M, Mort E.J, Desforges M, Moghimi A, Bartels B, El-Bacha T, Fowden A.L, Camm E.J. Diet-induced Maternal Obesity Impacts Feto-placental Growth and Induces Sex-specific Alterations in Placental Morphology, Mitochondrial Bioenergetics, Dynamics, Lipid Metabolism and Oxidative Stress in Mice. Acta Physiol. 2022;234:e13795.
- Powell T.L, Barner K, Madi L, Armstrong M, Manke J, Uhlson C, Jansson T, Ferchaud-Roucher V. Sex-Specific Responses in Placental Fatty Acid Oxidation, Esterification and Transfer Capacity to Maternal Obesity. Biochim. Biophys. Acta BBA Mol. Cell Biol. Lipids 2021;1866:158861.
- Bucher M, Montaniel K.R.C, Myatt L, Weintraub S, Tavori H, Maloyan A. Dyslipidemia, Insulin Resistance, and Impairment of Placental Metabolism in the Offspring of Obese Mothers. J. Dev. Orig. Health Dis. 2021;12:738–747.
- Jarvie E, Hauguel-de-Mouzon S, Nelson S.M, Sattar N, Catalano P.M, Freeman D.J. Lipotoxicity in Obese Pregnancy and Its Potential Role in Adverse Pregnancy Outcome and Obesity in the Offspring. Clin. Sci. 2010;119:123–129.
- Saben J, Lindsey F, Zhong Y, Thakali K, Badger T.M, Andres A, Gomez-Acevedo H, Shankar K. Maternal Obesity Is Associated with a Lipotoxic Placental Environment. Placenta 2014;35:171–177.
- Stewart F.M, Freeman D.J, Ramsay J.E, Greer I.A, Caslake M, Ferrell W.R. Longitudinal Assessment of Maternal Endothelial Function and Markers of Inflammation and Placental Function throughout Pregnancy in Lean and Obese Mothers. J. Clin. Endocrinol. Metab. 2007;92:969–975.
- Challier J.C, Basu S, Bintein T, Minium J, Hotmire K, Catalano P.M, Hauguel-de Mouzon S. Obesity in Pregnancy Stimulates Macrophage Accumulation and Inflammation in the Placenta. Placenta 2008;29:274–281.
- Roberts K.A, Riley S.C, Reynolds R.M, Barr S, Evans M, Statham A, Hor K, Jabbour H.N, Norman J.E, Denison F.C. Placental Structure and Inflammation in Pregnancies Associated with Obesity. Placenta 2011;32:247–254.
- Aye I.L.M.H, Lager S, Ramirez V.I, Gaccioli F, Dudley D.J, Jansson T, Powell T.L. Increasing Maternal Body Mass Index Is Associated with Systemic Inflammation in the Mother and the Activation of Distinct Placental Inflammatory Pathways1. Biol. Reprod. 2014;90:129.
- Zhou Y, Xu T, Wu Y, Wei H, Peng J. Oxidative Stress and Inflammation in Sows with Excess Backfat: Up-Regulated Cytokine Expression and Elevated Oxidative Stress Biomarkers in Placenta. Animals 2019;9:796.
- Pietro L, Daher S, Rudge M.V.C, Calderon I.M.P, Damasceno D.C, Sinzato Y.K, Bandeira C, Bevilacqua E. Vascular Endothelial Growth Factor (VEGF) and VEGF-Receptor Expression in Placenta of Hyperglycemic Pregnant Women. Placenta 2010;31:770–780.
- Pazinato F.M, Curcio B.D.R, Fernandes C.G, Santos C.A, Feijó L.S, Varela A.S, Nogueira C.E.W. Histomorphometry of the Placental Vasculature and Microcotyledons in Thoroughbred Mares with Chronic Laminitis. Theriogenology 2017;91:77–81.
- Sessions-Bresnahan D.R, Heuberger A.L, Carnevale E.M. Obesity in Mares Promotes Uterine Inflammation and Alters Embryo Lipid Fingerprints and Homeostasis. Biol. Reprod. 2018;99:761–772.
- Dunbar B, Bauer J. Metabolism of Dietary Essential Fatty Acids and Their Conversion to Long-Chain Polyunsaturated Metabolites. J. Am. Vet. Med. Assoc. 2002;220:1621–1626.
- King S, AbuGhazaleh A, Webel S, Jones K. Circulating Fatty Acid Profiles in Response to Three Levels of Dietary Omega-3 Fatty Acid Supplementation in Horses. J. Anim. Sci. 2007;86:1114–1123.
- Hodge L.B, Rude B.J, Dinh T.N, Lemley C.O. Effect of ω-3 Fatty Acid Supplementation to Gestating and Lactating Mares: On Milk IgG, Mare and Foal Blood Concentrations of IgG, Insulin and Glucose, Placental Efficiency, and Fatty Acid Composition of Milk and Serum from Mares and Foals. J. Equine Vet. Sci. 2017;51:70–78.
- . Principles of Animal Nutrition. CRC Press; Boca Raton, FL, USA 2018. Chapter 6: Metabolism and Nutrition of Lipids; p. 800.
- Islam A, Kodama T, Yamamoto Y, Ebrahimi M, Miyazaki H, Yasumoto Y, Kagawa Y, Sawada T, Owada Y, Tokuda N. Omega-3 Fatty Acids Transport through the Placenta. Asian J. Med. Biol. Res. 2016;2:1–8.
- Ortega-Senovilla H, Alvino G, Taricco E, Cetin I, Herrera E. Gestational Diabetes Mellitus Upsets the Proportion of Fatty Acids in Umbilical Arterial but Not Venous Plasma. Diabetes Care 2009;32:120–122.
- Elzinga S, Wood P, Adams A.A. Plasma Lipidomic and Inflammatory Cytokine Profiles of Horses with Equine Metabolic Syndrome. J. Equine Vet. Sci. 2016;40:49–55.
- O’Connor C.I, Lawrence L.M, Hayes S.H. Dietary Fish Oil Supplementation Affects Serum Fatty Acid Concentrations in Horses1. J. Anim. Sci. 2007;85:2183–2189.
- Dervishi E, Joy M, Sanz A, Alvarez-Rodriguez J, Molino F, Calvo J.H. Forage Preservation (Grazing vs. Hay) Fed to Ewes Affects the Fatty Acid Profile of Milk and CPT1B Gene Expression in the Sheep Mammary Gland. BMC Vet. Res. 2012;8:106.
- Pietrzak-Fiećko R, Kamelska-Sadowska A.M. The Comparison of Nutritional Value of Human Milk with Other Mammals’ Milk. Nutrients 2020;12:1404.
- Zentek J, Buchheit-Renko S, Ferrara F, Vahjen W, Van Kessel A.G, Pieper R. Nutritional and Physiological Role of Medium-Chain Triglycerides and Medium-Chain Fatty Acids in Piglets. Anim. Health Res. Rev. 2011;12:83–93.
- Takase S, Goda T. Effects of Medium-Chain Triglycerides on Brush Border Membrane-Bound Enzyme Activity in Rat Small Intestine. J. Nutr. 1990;120:969–976.
- Dierick N.A, Decuypere J.A, Molly K, Van Beek E, Vanderbeke E. The Combined Use of Triacylglycerols (TAGs) Containing Medium Chain Fatty Acids (MCFAs) and Exogenous Lipolytic Enzymes as an Alternative to Nutritional Antibiotics in Piglet Nutrition. Livest. Prod. Sci. 2002;76:1–16.
- Kono H, Fujii H, Asakawa M, Maki A, Amemiya H, Hirai Y, Matsuda M, Yamamoto M. Medium-Chain Triglycerides Enhance Secretory IgA Expression in Rat Intestine after Administration of Endotoxin. Am. J. Physiol.-Gastrointest. Liver Physiol. 2004;286:G1081–G1089.
- Wang J, Wu X, Simonavicius N, Tian H, Ling L. Medium-Chain Fatty Acids as Ligands for Orphan G Protein-Coupled Receptor GPR84. J. Biol. Chem. 2006;281:34457–34464.
- de la Garza Puentes A, Martí Alemany A, Chisaguano A.M, Montes Goyanes R, Castellote A.I, Torres-Espínola F.J, García-Valdés L, Escudero-Marín M, Segura M.T, Campoy C. The Effect of Maternal Obesity on Breast Milk Fatty Acids and Its Association with Infant Growth and Cognition—The PREOBE Follow-Up. Nutrients 2019;11:2154.
- Armand M, Bernard J.Y, Forhan A, Heude B, Charles M.-A, Annesi-Maesano I, Bernard J.Y, Botton J, Charles M.-A, Dargent-Molina P. Maternal Nutritional Determinants of Colostrum Fatty Acids in the EDEN Mother-Child Cohort. Clin. Nutr. 2018;37:2127–2136.
- Rydlewski A.A, Manin L.P, Pizzo J.S, Silva P.D, da Silveira R, Tavares C.B.G, de Paula M, Pereira O, Santos O.O, Visentainer J.V. Lipid Profile by Direct Infusion ESI-MS and Fatty Acid Composition by GC-FID in Human Milk: Association with Nutritional Status of Donors. J. Food Compos. Anal. 2021;100:103797.
- Tang J. Advances in Dietary Lipids and Human Health. Elsevier; Amsterdam, The Netherlands 2022. Palmitoleic Acid in Health and Disease; pp. 293–302.
- Duckett S.K, Volpi Lagreca G, Alende M, Long N. Palmitoleic Acid Reduces Intramuscular Lipid and Restores Insulin Sensitivity in Obese Sheep. Diabetes Metab. Syndr. Obes. Targets Ther. 2014;7:553–563.
- de Souza C.O, Teixeira A.A.S, Biondo L.A, Lima Junior E.A, Batatinha H.A.P, Rosa Neto J.C. Palmitoleic Acid Improves Metabolic Functions in Fatty Liver by PPARα-Dependent AMPK Activation: PALMITOLEATE CONTROLLING PPARα-AMPK ON LIVER. J. Cell. Physiol. 2017;232:2168–2177.
- Yao D, Luo J, He Q, Shi H, Li J, Wang H, Xu H, Chen Z, Yi Y, Loor J.J. SCD1 Alters Long-Chain Fatty Acid (LCFA) Composition and Its Expression Is Directly Regulated by SREBP-1 and PPARγ 1 in Dairy Goat Mammary Cells: SCD1 AND FATTY ACID METABOLISM. J. Cell. Physiol. 2017;232:635–649.
- di Martino O, Troiano A, Addi L, Guarino A, Calabrò S, Tudisco R, Murru N, Cutrignelli M.I, Infascelli F, Calabrò V. Regulation of Stearoyl Coenzyme A Desaturase 1 Gene Promoter in Bovine Mammary Cells. Anim. Biotechnol. 2015;26:251–259.
- Dobrzyn P, Jazurek M, Dobrzyn A. Stearoyl-CoA Desaturase and Insulin Signaling—What Is the Molecular Switch?. Biochim. Biophys. Acta BBA Bioenerg. 2010;1797:1189–1194.
- Shi H.B, Zhao W.S, Luo J, Yao D.W, Sun Y.T, Li J, Shi H.P, Loor J.J. Peroxisome Proliferator-Activated Receptor Γ1 and Γ2 Isoforms Alter Lipogenic Gene Networks in Goat Mammary Epithelial Cells to Different Extents. J. Dairy Sci. 2014;97:5437–5447.
- Flint D.J, Travers M.T, Barber M.C, Binart N, Kelly P.A. Diet-Induced Obesity Impairs Mammary Development and Lactogenesis in Murine Mammary Gland. Am. J. Physiol.-Endocrinol. Metab. 2005;288:E1179–E1187.
- Marko R.C, Branislava B, Bojan T, Aleksandar P, Milenko S, Branislav L, Ivan R. Metabolic Acclimation to Heat Stress in Farm Housed Holstein Cows with Different Body Condition Scores. Afr. J. Biotechnol. 2011;10:10293–10303.
- Liu Z, Ezernieks V, Wang J, Arachchillage N.W, Garner J.B, Wales W.J, Cocks B.G, Rochfort S. Heat Stress in Dairy Cattle Alters Lipid Composition of Milk. Sci. Rep. 2017;7:961.
- Bolzan A.M.S, Bonnet O.J.F, Wallau M.O, Basso C, Neves A.P, Carvalho P.C.d.F. Foraging Behavior Development of Foals in Natural Grassland. Rangel. Ecol. Manag. 2020;73:243–251.
- Contu L, Hawkes C. A Review of the Impact of Maternal Obesity on the Cognitive Function and Mental Health of the Offspring. Int. J. Mol. Sci. 2017;18:1093.
- Thompson J.R, Valleau J.C, Barling A.N, Franco J.G, De Capo M, Bagley J.L, Sullivan E.L. Exposure to a High-Fat Diet during Early Development Programs Behavior and Impairs the Central Serotonergic System in Juvenile Non-Human Primates. Front. Endocrinol. 2017;8:164.
- Muhlhausler B, Adam C, Findlay P, Duffield J, McMillen I. Increased Maternal Nutrition Alters Development of the Appetite-Regulating Network in the Brain. FASEB J. 2006;20:1257–1259.
- Purcell R, Sun B, Pass L, Power M, Moran T, Tamashiro K. Maternal Stress and High-Fat Diet Effect on Maternal Behavior, Milk Composition, and Pup Ingestive Behavior. Physiol. Behav. 2011;104:474–479.
- Borengasser S.J, Lau F, Kang P, Blackburn M.L, Ronis M.J.J, Badger T.M, Shankar K. Maternal Obesity during Gestation Impairs Fatty Acid Oxidation and Mitochondrial SIRT3 Expression in Rat Offspring at Weaning. PLoS ONE 2011;6:e24068.
- Isganaitis E, Woo M, Ma H, Chen M, Kong W, Lytras A, Sales V, De Coste-Lopez J, Lee K.-J, Leatherwood C. Developmental Programming by Maternal Insulin Resistance: Hyperinsulinemia, Glucose Intolerance, and Dysregulated Lipid Metabolism in Male Offspring of Insulin-Resistant Mice. Diabetes 2014;63:688–700.
- Boyle K.E, Patinkin Z.W, Shapiro A.L.B, Bader C, Vanderlinden L, Kechris K, Janssen R.C, Ford R.J, Smith B.K, Steinberg G.R. Maternal Obesity Alters Fatty Acid Oxidation, AMPK Activity, and Associated DNA Methylation in Mesenchymal Stem Cells from Human Infants. Mol. Metab. 2017;6:1503–1516.
- Musaev A, Sadykova S, Anambayeva A, Saizhanova M, Balkanay G, Kolbaev M. Mare’s Milk: Composition, Properties, and Application in Medicine. Arch. Razi Inst. 2021;76:1125–1135.
- . State of the Climate in Europe 2022. WMO 2023. p. 40.
Use Nutrition Calculator
Check if your horse's diet meets their nutrition requirements with our easy-to-use tool Check your horse's diet with our easy-to-use tool
Talk to a Nutritionist
Discuss your horse's feeding plan with our experts over a free phone consultation Discuss your horse's diet over a phone consultation
Submit Diet Evaluation
Get a customized feeding plan for your horse formulated by our equine nutritionists Get a custom feeding plan formulated by our nutritionists