Effect of 25-hydroxyvitamin D3 on periparturient Yili mare and foal health

The periparturient period in mares is metabolically demanding and requires optimal nutrition to support maternal health and fetal development. Vitamin D₃, particularly its metabolite 25(OH)D₃, plays crucial roles beyond bone health, including immune and antioxidant functions. This study investigated the effects of supplementing pregnant and lactating mares with 25-hydroxyvitamin D₃ (25(OH)D₃) at a dose of 120,000 IU/day during late gestation and early lactation. Twelve Yili horses were randomly divided into two groups of six, matched by gestation and foaling date. One group received a standard diet, while the other was supplemented with 120,000 IU/day of 25(OH)D₃ for 60 days (30 days before and 30 days after birth). Serum and milk samples were collected at the start and midpoint (days 0 and 30) for analysis of 25(OH)D₃, mineral concentrations (calcium and phosphorus), proteins, lipids, antioxidants, immunoglobulins, and cytokine levels. The supplementation significantly elevated serum and milk concentrations of 25(OH)D₃, calcium, phosphorus, glucose, total protein, and globulin, whereas bilirubin levels diminished in both mares and foals (P < 0.05). In supplemented mares, milk fat percentage, protein content, and immunoglobulin levels (IgA, IgG, and IgM) were elevated (P < 0.05). Improvements were observed in immunoglobulin levels, inflammatory markers, and antioxidant capacity in the supplemented group (P < 0.05). In conclusion, 25(OH)D₃ supplementation in pregnant and lactating mares enhanced serum and milk nutrient levels, decreased bilirubin levels, and increased immunoglobulin levels. It also boosted antioxidant capacity and immunoglobulin levels and reduced cytokine levels in both mares and foals.

1 Introduction

The periparturient period is a critical physiological stage in broodmares that spans from late gestation to the initial weeks postpartum. During this period, mares undergo complex physiological adaptations associated with pregnancy, parturition, and subsequent lactation onset (Del Prete et al., 2024; Hassan et al., 2025). This period is typically divided into two distinct phases: the prepartum period (30 days before foaling) and the postpartum period (30 days after foaling). Due to the intricate physiological changes that occur during this period, the maternal system becomes particularly vulnerable to metabolic stress, making it a high-risk period for the development of various metabolic disorders (Jung et al., 2021; Purru Kanakaiah et al., 2023). Maternal metabolism is crucial for nutrient transfer to the placenta, which in turn affects fetal growth (Nogues et al., 2021). Recent advancements in the mother-offspring integration concept have prompted increased research on targeted nutritional and metabolic interventions to optimize reproductive performance in animals (Simon et al., 2015; Kamal et al., 2023a; Kamal et al., 2023c). This integrative approach enables the strategic modulation of maternal-offspring nutritional crosstalk, facilitating postpartum recovery in broodmares, enhancing their subsequent reproductive capacity, and ensuring robust neonatal development (Connysson and Saastamoinen, 2023). Maternal factors directly influence the physiological status and production performance of offspring, a phenomenon known as maternal effects. Notably, the peripartum nutritional and metabolic status of dams exerts profound regulatory effects on offspring physiology and production performance (Herring et al., 2018; Connysson and Saastamoinen, 2023).

Vitamin D₃ (cholecalciferol), an essential micronutrient in livestock production, plays critical functions beyond its classical role in calcium-phosphorus homeostasis and rickets prevention. Emerging evidence highlights its pleiotropic effects, including anti-inflammatory, antioxidant, and immune-enhancing properties, in domestic animals (Upadhaya et al., 2021; Babić Leko et al., 2023). Studies have demonstrated that maternal vitamin D₃ deficiency during gestation is associated with an elevated risk of preterm birth, impaired fetal skeletal development, osteomalacia, and compromised production performance (Zhang and Piao, 2019; Biehler-Gomez et al., 2024). Additionally, Del Prete et al (Del Prete et al., 2024). discovered that dietary supplementation with Oxyliver^® (a commercial formulation comprising a blend of antioxidants) from 290 to 320 days of gestation in Italian Salernitano mares correlates with a decreased gestation duration and an enhancement in liver functions, ultimately improving colostrum quality. The need for calcium and phosphorus in pregnant mares markedly escalates throughout the final trimester to facilitate embryonic skeletal development (Upadhaya et al., 2021). Vitamin D is crucial for the absorption of these elements. Nevertheless, well-managed horses fed high-quality fodder with consistent turnout generally maintain sufficient vitamin D levels during gestation. Despite sufficient vitamin D levels, supplementation of mares with 25(OH)D₃ could improve the quality of colostrum (Upadhaya et al., 2021; Connysson and Saastamoinen, 2023). 25(OH)D₃ shows greater bioavailability and reduced hepatic processing than vitamin D₃, enhancing its supplement efficacy (Cashman et al., 2012; Jetter et al., 2014; Christakos et al., 2016).

Supplementation of periparturient dairy cows with 120,000 IU/d of 25(OH)D₃ enhanced the immune function of both the cow and calf, and this amount also increased the quality of colostrum and antioxidant-immune profiles (Xu et al., 2021a). This dosage has been reported to enhance colostrum quality and improve the antioxidant and immune profiles of both dams and calves (Ahmadi and Mohri, 2021; Xu et al., 2021a). Serum 25(OH)D₃ levels in pasture mares and foals remain unknown under grazing conditions, which may negatively affect their productive performance (Alemi et al., 2025). There is limited information regarding the impact of 25(OH)D₃ on the antioxidant balance and immune system of mares and foals. Considering that the feeding dynamics and time-activity budgets of grazing horses—which significantly influence nutrient intake and metabolic state—can vary with management systems (Lamanna et al., 2025), targeted nutritional strategies are crucial. Therefore, administering 120,000 IU/day of D₃ (25 (OH) D₃) to pregnant and nursing mares during late pregnancy and early lactation may improve the health of both the mother and foal by improving the metabolism of calcium and phosphorus, the immune system, and the ability to fight off free radicals. This treatment is expected to improve the immune health of both mares and foals by increasing cytokine regulation and immunoglobulin levels.

2 Materials and methods

2.1 Ethical approval

All experimental procedures involving animals were approved (animal protocol number: 2024019) by the Animal Welfare and Ethics Committee of Xinjiang Agricultural University, Urumqi, Xinjiang, China.

2.2 Animals, diets, and experimental design

This experiment was conducted at Zhaosu Stud Farm (Yili, Xinjiang, China) from May to July 2024. The study utilized twelve multiparous Yili mares, six mares in each group, aged 5–7 years, during the peripartum period. To reduce variability, the mares were grouped based on the number of times they had given birth and when they were expected to give birth. All mares were fed a concentrate supplement (Table 1). Within each block, mares were randomly assigned to one of two treatments in a randomized block design (Del Prete et al., 2024): Control group: each mare received a daily concentrate supplement of 500 g without additional supplementation (Hassan et al., 2025); Treatment group: each mare received a daily concentrate supplement of 500 g, supplemented with 6 g of a 25(OH)D₃ product provided by Jiangsu Caiwei Biotechnology Co., Ltd. (Nanjing, China) and contained 20,000 IU/g of 25(OH)D₃. The active ingredient, 25-hydroxyvitamin D₃, was uniformly diluted in a carrier of starch and dextrin. The declared concentration of 25(OH)D₃ in the final product was 0.05% (500 ppm). The precise daily dose of 25(OH)D₃ for each mare was first placed on the surface of a small portion of the concentrate supplement in the feeding bucket. The foals were allowed to consume this initial portion completely under supervision before the remainder of their daily concentrate ration was provided. This two-step feeding strategy was designed specifically to ensure that the supplement was consumed in its entirety before the main feed, thereby minimizing the risk of loss or selective refusal.

Table 1

Table 1. Ingredients and chemical composition of the concentrate supplement (dry matter basis).

The prepartum supplementation period averaged 30 days (range: 28–32 days). Postpartum, mares continued to receive the same dietary treatment (including 25(OH)D₃ supplementation) for an additional 30 days, resulting in a total supplementation period of 60 days. Throughout the trial period, the mares were provided with free access to clean water and were maintained under a grazing regime with supplemental feeding. After birth, foals were allowed to suckle freely from their dams.

2.3 Sampling

The first day of the experiment was designated as -30 d (relative to foaling). Blood sampling was performed before morning feeding on two consecutive days (days 0 and 30). Jugular venous blood was drawn from all mares and their post-colostrum foals using standard vacuum blood collection tubes. Blood samples in plain tubes were allowed to clot for 30 min at room temperature, followed by centrifugation at 2,000 × g for 15 min. Serum aliquots were transferred to 1.7-mL cryovials and stored at -20°C until analysis. Serum 25(OH)D₃ was quantified utilizing a commercial ELISA kit (Elaescience Bioscience and Technology Co., Ltd., Wuhan, China).

2.4 Biochemical analysis

Metabolic parameters including total protein (TP), albumin (ALB), globulin (GLB), glucose (Glu), triglyceride (TG), total cholesterol (TC), high-density lipoprotein (HDL), low-density lipoprotein (LDL), total bilirubin (TB), calcium (Ca), and phosphorus (P) were analyzed using an automated biochemical analyzer (BS-240VET, Shenzhen Myriad Biomedical Co., Ltd., China).

2.5 Antioxidant and immunity activity

Superoxide dismutase (SOD), total antioxidant capacity (T-AOC), catalase (CAT), and glutathione peroxidase (GSH-Px) were measured using colorimetric kits. Malondialdehyde (MDA) levels were determined via the thiobarbituric acid-reactive substances assay. All antioxidant-related kits were obtained from Nanjing Jiancheng Institute of Bioengineering (Nanjing, China). Immunoglobulin (IgA, IgG, and IgM) and cytokine (IL-1β, IL-2, IL-6, IL-10, and TNF-α) concentrations were analyzed using ELISA, according to Wang et al. (Wang et al., 2022). Horse-specific ELISA kits were purchased from Nanjing Jiancheng Institute of Bioengineering. Serum testing was not diluted.

2.6 Fat, protein, lactose, and minerals detection

Mare milk was collected by artificial milking on two consecutive sampling days (postpartum days 0 and 30). The milk was not treated in any way before the tests. For each mare, a 24-hour composite sample (50 mL) was created by volume-weight mixing of three milk collections, adjusted according to production levels. Mares’ milk specimens were stored at 4°C and analyzed using the MilkoScan™ FT3 Dairy Analyzer (FOSS, Model 91889105) for fat, protein, lactose, and Solids-not-fat (SNF). The calcium content was determined to analyze the mare’s milk concentration using the EDTA Back-Titration Method. Phosphorus quantification employed ammonium vanadate-molybdate spectrophotometric detection at 420 nm. A horse 25(OH)D₃ ELISA (El-aescience Bioscience and Technology Corporation, Wuhan, China) was used to determine mare’s milk 25(OH)D₃.

2.7 Statistical analysis

Data was analyzed using SPSS 19.0 software (SPSS 19.0, IBM Corporation, Armonk, NY, USA). Data were analyzed using a repeated measures analysis of variance (ANOVA) model. Duncan’s method of multiple comparisons was used to identify significant differences (P ≤ 0.05) between treatments. Before analysis, assumptions of normality (assessed via the Shapiro-Wilk test) and homogeneity of differences (evaluated using Levene’s test) were verified. The normality of all data distributions was confirmed using the Shapiro-Wilk test (P > 0.05), justifying the use of parametric statistical methods.

3 Results

3.1 Effects of 25(OH)D₃ supplementation on calcium and phosphorus concentrations

Table 2 illustrates that the supplemented mares demonstrated significantly elevated serum 25(OH)D₃, calcium (Ca), and phosphorus (P) levels on day 0 (P < 0.05 and P < 0.01). Although mares maintained notably elevated serum 25(OH)D₃ levels (P < 0.01), calcium concentrations returned to baseline by day 30 compared to non-supplemented mares.

Table 2

Table 2. Effects of dietary 25(OH)D₃ supplementation on serum 25(OH)D₃, calcium, and phosphorus concentrations in mares and foals.

Additionally, Table 2 also illustrates that foals from supplemented dams displayed significantly elevated serum 25(OH)D₃ levels (210.91%, P < 0.01), calcium (18.92%, P < 0.05), and phosphorus (27.16%, P < 0.05) on day 0. Additionally, supplemented foals maintained significantly higher serum 25(OH)D₃ levels (48.96% increase, P < 0.01), while calcium and phosphorus values on day 30 were comparable to those of non-supplemented mares.

3.2 Effects of 25(OH)D₃ supplementation on milk composition, and minerals

Table 3 demonstrates that the supplemented mares showed marked increases in milk protein (36.22%, P < 0.01) and 25(OH)D₃ (572.32%, P < 0.01); additionally, milk fat, calcium, and phosphorus levels were elevated (P < 0.05) on day 0. At day 30, milk from supplemented mares continued to show numerically higher concentrations of fat, protein, calcium, phosphorus, and solids-not-fat compared to the control; however, these differences were not statistically significant (P > 0.05). In contrast, the concentration of 25(OH)D₃ in milk remained substantially elevated in the supplemented group (506.90%, P < 0.01).

Table 3

Table 3. Effects of dietary 25(OH)D₃ supplementation on mare’s milk composition, 25(OH)D₃, calcium, and phosphorus concentrations.

3.3 Effects of dietary 25(OH)D₃ supplementation on serum indicators in mares and foals

Table 4 shows that mares receiving the supplements had significantly higher serum concentrations of total protein (TP), globulin, and triglycerides, and serum glucose levels increased by 4.7%, while total bilirubin (TB) levels decreased by 9.7%, although these changes were not statistically significant (P > 0.05) on day 0. The results indicated that mares receiving the supplements had higher serum concentrations of TP, globulin, and glucose (P < 0.01), whereas TB levels were markedly lower (P < 0.01) on day 30 than in mares not receiving the supplements.

Table 4

Table 4. Effects of dietary 25(OH)D₃ supplementation on serum indices in mares and foals.

Furthermore, foals from mares receiving 25(OH)D₃ supplementation showed significant increases in serum TP (P < 0.01), globulin (P < 0.01), and triglyceride levels (P < 0.01) on day 0. The results indicated that foals of mares receiving 25(OH)D₃ supplementation had notably increased serum TP concentrations (7.46%, P < 0.05) and globulin levels (43.98%, P < 0.01), while TB (P < 0.01), albumin (P < 0.01), and glucose levels (P < 0.05) were decreased on day 30 compared to foals of mares that did not receive the supplements.

3.4 Impacts of dietary 25(OH)D₃ supplementation on serum antioxidant in mares and foals

Figure 1 and Supplementary 1 show that mares receiving the supplements had substantially higher serum SOD and GSH-Px activities (P < 0.01). Meanwhile, MDA levels were significantly lower (P < 0.01), and there was a significant increase in serum SOD activity (P < 0.01) on day 0. The results indicated that in mares receiving the supplements, SOD and GSH-Px activity remained highly elevated (P < 0.01), CAT and T-AOC activity were significantly increased (P < 0.05), and MDA levels were markedly decreased (P < 0.05) on day 30. Furthermore, foals receiving the supplements showed significantly higher GSH-Px activity (P < 0.01), along with significant increases in CAT activity (P < 0.05) compared to mares that did not receive the supplements.

Figure 1

Figure 1. Effects of dietary 25(OH)D₃ supplementation on antioxidant function in mares and foals. SOD, superoxide dismutase; CAT, catalase; GSH-Px, glutathione peroxidase; MDA, malondialdehyde; T-AOC, total antioxidant capacity. Significance level stated the values, *P < 0.05, **P < 0.01.

By day 30 postpartum (mares), serum SOD and GSH-Px activities were substantially higher (P < 0.01) in the supplemented group than in the control. Concurrently, CAT activity and T-AOC increased notably (P < 0.05), whereas MDA levels decreased (P < 0.05). At 30 days postpartum, supplemented foals exhibited elevated GSH-Px activity (P < 0.01). Moreover, CAT activity was significantly increased (P < 0.05).

3.5 Effect of dietary 25(OH)D₃ supplementation on serum and milk immunoglobulin in mares and foals

Figure 2 and Supplementary 2 demonstrate that mares receiving the supplements showed significant increases in serum IgA, IgG, and IgM levels (P < 0.01). The concentrations of IgA, IgG, and IgM in mare milk were also markedly increased (P < 0.01). In foal serum, IgG levels were substantially increased (P < 0.01), with a significant increase in IgA levels (P < 0.05) on postpartum day 0. The results indicated that supplemented mares still had significantly elevated mare serum IgA, IgG, and IgM levels (P < 0.01). The IgG concentration in mare milk was also notably higher (P < 0.01). Supplemented foals showed highly significant increases in both IgG and IgA (P < 0.01), with marginal significance as well for IgA (P < 0.05) on day 30 compared to mares that did not receive the supplement.

Figure 2

Figure 2. Effects of dietary 25(OH)D₃ supplementation on immune function in mares and foals. IgA, immunoglobulin A; IgM, immunoglobulin M; IgG, immunoglobulin G. Significance level stated the values, *P < 0.05, **P < 0.01.

3.6 Effects of dietary 25(OH)D₃ addition on serum inflammatory cytokines in mares and foals

Figure 3 and Supplementary 3 illustrate the effects of dietary 25(OH)D₃ supplementation on serum inflammatory cytokine levels in mares and foals. On the first day after giving birth, the mares that received the supplement had much lower levels of IL-1β (P < 0.05) and showed substantial decreases in IL-2, IL-6, TNF-α, and IL-10 compared to those that did not receive the supplement (P < 0.01). Serum-supplemented foals exhibited highly significant reductions in IL-1β, IL-2, IL-6, and TNF-α levels (P < 0.01).

Figure 3

Figure 3. Influence of dietary 25(OH)D₃ supplementation on serum inflammatory cytokines in mares and foals. IL-1β, interleukin-1β; IL-2, interleukin-2; IL-6, interleukin-6; IL-10, interleukin-10; TNF-α, tumor necrosis factor-α. Significance level stated the values, *P < 0.05, **P < 0.01.

By day 30 postpartum (mares’ serum), IL-1β, IL-2, IL-6, TNF-α, and IL-10 levels remained markedly lower in supplemented groups (P < 0.01). On day 30, the supplemented group demonstrated significant reductions in IL-2, IL-6, and TNF-α (P < 0.01) compared to the control.

4 Discussion

Maternal nutritional management during gestation and lactation plays an important role in maintaining maternal health and fostering offspring growth and immune system development. Our results are consistent with the results of previous experimental studies, which showed that supplementing periparturient dairy cows with 120,000 IU/d of 25(OH)D₃ notably boosts serum concentrations of the vitamins and enhances antioxidant capacity and immune response in dams and calves (Alhussien et al., 2021; Somagond et al., 2023; Yadav et al., 2024). The transfer of 25(OH)D₃ to the fetus occurs via active placental endocytosis with a 27% uptake efficiency, highlighting the essential role of the placenta in maternal-fetal nutrient transfer (Ashley et al., 2022). The predominant form of vitamin D in maternal milk is 25(OH)D₃, showing a strong association between milk and maternal serum levels of this metabolite (Jan Mohamed et al., 2014; Yu et al., 2021). Maternal vitamin D deficiency reduces vitamin D levels in milk, thereby increasing the vulnerability of offspring to deficiencies (Schossow et al., 2018). Additionally, Azarpeykan et al (Azarpeykan et al., 2022). found that horses make negligible quantities of vitamin D₃ in their skin after exposure to UVB light and may therefore rely on their diet as a primary source of vitamin D. Serum 25(OH)D₃ levels in grazing mares and foals remain suboptimal under pasture conditions, potentially negatively affecting their production performance (Mainguy-Seers et al., 2024; Alemi et al., 2025). For mares, detailed studies on the placental absorption efficiency and precise mechanism of 25(OH)D₃ have not yet been conducted. However, this study finds that the concentration of 25(OH)D₃ in the serum of foals significantly increases after parturition, indicating that 25(OH)D₃ can be transferred to foals via the placenta and demonstrating that similar mechanisms may potentially exist.

Vitamin D metabolic processes are critical for maintaining mineral homeostasis, particularly regulating calcium-phosphorus equilibrium (Cândido and Bressan, 2014; Burns-Whitmore et al., 2024). The primary vitamin D metabolite, 25(OH)D₃, is the gold standard biomarker for evaluating vitamin D status, as its plasma concentration directly reflects systemic availability (Tuckey et al., 2019; Alonso et al., 2023). The biologically active 1,25(OH)2D₃ promotes calcium absorption in the intestine and maintains skeletal integrity through VDR-dependent genomic signaling in enterocytes (Haussler et al., 2013; Fleet, 2022; Voltan et al., 2023). Maintaining adequate 25(OH)D₃ levels is indispensable for preserving calcium-phosphorus homeostasis and facilitating proper growth in animal populations. Experimental supplementation with 25(OH)D₃ in periparturient mares led to notable increases in maternal and offspring serum concentrations of calcium, phosphorus, and 25(OH)D₃. The observed outcomes mirror those of Xu et al (Xu et al., 2021a). in dairy cows, who found that 25(OH)D₃ supplementation during the periparturient period significantly boosted plasma levels of total calcium, phosphorus, and vitamin D metabolites within three weeks postpartum.

The percentage of protein and fat content in milk is an essential quality parameter for evaluating its composition. 25(OH)D₃ modulates mammary immune competence, mitigates mastitis symptoms, and increases expression of genes linked to milk protein synthesis, such as CDL62 and iNOS (Poindexter et al., 2020). Additionally, Martinez et al (Martinez et al., 2018). showed that prenatal administration of 25 (OH) D₃ resulted in higher colostrum protein, lactose, and total solid concentrations. Furthermore, sows that received dietary 25(OH)D₃ supplementation had elevated levels of milk protein and lactose in their milk (Zhou et al., 2017).

Our findings indicate that 25(OH)D₃ supplementation substantially increased the protein and fat content of colostrum. Xu et al. (Xu et al., 2021a) demonstrated that prepartum provision of 25(OH)D₃ substantially elevated Ca, P, and 25(OH)D₃ concentrations in colostrum and mature milk within 21 days postpartum. This study also showed that administering 25(OH)D_3–30 days before parturition significantly increased the protein and fat content in colostrum and elevated the levels of calcium, phosphorus, and 25(OH)D₃ in both colostrum and mature milk collected 30 days postpartum.

Circulating biochemical parameters are key diagnostic tools for monitoring systemic metabolic equilibrium and physiological well-being in livestock species (Kamal et al., 2023b). The albumin and globulin fractions in serum mediate three core physiological functions: plasma oncotic pressure maintenance, bioactive compound transportation, and immunological response regulation (Levitt and Levitt, 2016). The serum total protein-enhancing effect of 120,000 IU/d 25(OH)D₃ supplementation observed in cows (Silva et al., 2022) was replicated in the present equine model. Likewise, dietary 25(OH)D₃ supplementation markedly elevated serum TP and globulin levels in mares and foals under trial conditions. These findings suggest a potential association between the observed effects and immunomodulatory functions of 25(OH)D₃. Increased total bilirubin concentrations are diagnostic indicators of severe hepatic lipidosis in dairy cows (Sejersen et al., 2012). The characteristic reduction in hepatic detoxification capacity during the periparturient phase leads to systemic accumulation of bilirubin (Giannuzzi et al., 2021). Supplementation with 25(OH)D₃ decreased circulating bilirubin levels by 32–41% in mares and their offspring, indicating enhanced hepatocyte metabolism. Serum glucose and triglyceride levels are critical metabolites involved in bodily metabolism and play essential roles in maintaining energy homeostasis and physiological functions. The results demonstrated that 25(OH)D₃ supplementation notably elevated serum glucose concentrations in both mares and foals, while also increasing serum triglyceride levels in neonatal foals, indicating its regulatory role in energy metabolism.

The transition phase surrounding parturition is pivotal in broodmare management, where dietary interventions critically determine maternal-fetal health outcomes (Benyshek et al., 2006). Notably, the peripartum nutritional and metabolic status of dams exerts profound regulatory impacts on offspring physiology and production performance (Sejersen et al., 2012; Herring et al., 2018). 25(OH)D₃ enhances antioxidant defenses by stimulating SOD and GSH-Px biosynthesis, effectively scavenging ROS. In hyperglycemic human endothelial cells, 25(OH)D₃ reduces ROS/MDA levels and elevates GSH levels (Zhan et al., 2023). Similar 25(OH)D₃-induced increases in T-AOC, SOD, and GSH-Px and a decrease in MDA were observed in dairy cattle (Xu et al., 2021a). This study demonstrated that 25(OH)D₃ supplementation substantially increased serum SOD and GSH-Px levels and decreased MDA concentrations in both mares and foals. The treatment increased SOD and GSH-Px activities while decreasing MDA in dam-offspring pairs. These data establish 25(OH)D₃ as an effective supplement for optimizing redox balance in periparturient equine pairs.

The peripartum period in dairy cows is characterized by intense metabolic stress, negative energy balance, and inflammation, leading to a well-documented surge in ROS and oxidative stress markers such as MDA, alongside a depletion of endogenous antioxidants such as SOD and GSH-Px. This oxidative burden markedly contributes to immunosuppression and increases susceptibility to illnesses such as mastitis and metritis (Abuelo et al., 2019). The observation that 25(OH)D₃ supplementation increased SOD and GSH-Px levels while decreasing MDA levels is consistent with the established immunomodulatory and antioxidant functions of vitamin D metabolites. 25(OH)D₃, the principal circulating metabolite, is converted intracellularly to the active hormonal form, 1,25-dihydroxyvitamin D₃. Calcitriol (1,25-dihydroxycholecalciferol) functions through the vitamin D receptor, a nuclear receptor present in most immune cells and diverse tissues, to enhance the expression of genes associated with antioxidant defense mechanisms, such as those coding for SOD and GSH-Px, while possibly inhibiting pathways that produce excessive ROS or lipid peroxidation byproducts such as MDA (Jeong et al., 2024). Consequently, administering 25(OH)D₃ to periparturient dairy cows—considered more bioavailable and less susceptible to hydroxylation constraints than vitamin D₃ during metabolic stress—emerges as a promising nutritional approach to enhance antioxidant capacity, reduce oxidative damage (indicated by decreased MDA levels), and potentially enhance overall health and productivity during this critical period (Xu et al., 2021b).

Serum immunoglobulins are important for the immune response because they help recognize antigens and neutralize pathogens. These immunoproteins execute immunological surveillance through opsonization-mediated bacterial clearance mechanisms. Previous research has reported elevated serum IgG levels in transition dairy cows supplemented with 25(OH)D₃ (Xu et al., 2021a). Ruminant placentation restricts transplacental immunoglobulin transfer, requiring neonatal immunoglobulin intake through colostrum (Borghesi et al., 2014). Additionally, Martinez et al (Martinez et al., 2018). observed increased colostral IgG levels following 25 (OH) D₃ supplementation. Furthermore, Zhang et al (Zhang et al., 2019). found that administration of 25(OH)D₃ to pregnant pigs led to higher levels of IgG in their milk. 25(OH)D₃ supplementation increased the levels of IgA, IgG, and IgM in the blood of the mare, her milk, and the blood of the foal, which improved their immune system. Therefore, it can be noted that maternal immunoglobulin and 25(OH)D₃ concentrations are likely to be transferred to offspring through breastfeeding. Our findings align with those of Aoki et al. (Aoki et al., 2020), who established that prepartum 25(OH)D₃ supplementation in mares markedly elevated IgG concentrations in colostrum and serum IgG levels in foals, associated with diminished rates of passive transfer failure. Moreover, this corresponds with research on dairy cattle (Urakawa et al., 2024), indicating that a higher vitamin D status improved the quality of colostral immunoglobulins. In addition to passive transfer through immunoglobulins (with IgG being crucial for foals and calves), 25(OH)D₃ directly influences immune modulation in neonates. It operates through vitamin D receptors on immune cells, facilitating the synthesis of antimicrobial peptides, altering Tcell responses, and augmenting macrophage function, as shown by Ghaseminejad-Raeini et al. (Ghaseminejad-Raeini et al., 2023). Consequently, 25(OH)D₃ supplementation offers two advantages: enhancing immunoglobulin-rich colostrum production in the dam and directly bolstering the developing innate and adaptive immune systems of the neonate via transferred calcidiol (25-hydroxyvitamin D). Mares may assimilate 25(OH)D₃ more effectively than vitamin D₂ and D₃, rendering it a strategic supplement for periparturient immunity (Alemi et al., 2025).

Pro-inflammatory cytokines are likely to serve as essential indicators for assessing systemic immunocompetence. Inflammatory cytokines encompass diverse immune modulators participating in inflammatory cascades, with TNF-α, IL-1β, IL-2, and IL-6 representing the predominant pro-inflammatory subtypes (Borghesi et al., 2014). 25(OH)D₃ suppresses inflammatory cytokine production while improving immunocompetence in bovine dams and neonates (Xu et al., 2021a). 1, 25(OH)2D₃, the active metabolite of 25(OH)D₃, suppresses pro-inflammatory cytokine synthesis (IFN-γ/IL-17/IL-21), enhances CTLA-4 and FoxP3 Treg development, modulates Th1/Th2 polarization via CTLA-4-mediated pathways, and attenuates inflammatory responses through IL-10-dependent regulation (Zhang et al., 2018). Our findings revealed that 25(OH)D₃ supplementation reduced cytokine levels. This effect may occur through 25(OH)D3’s capacity to suppress IFN-γ-driven inflammation pathways that stimulate IL-1/IL-6/IL-8/TNF-α production (Baeke et al., 2010; Palmer et al., 2011). Vitamin D, crucial for periparturient mares and dairy cows, suppresses NF-κB and promotes anti-inflammatory Treg responses, improving uterine health and reducing inflammation postpartum (Guo et al., 2018). Although measuring cytokines indicates that the immune system is functioning, it does not provide a clear indication of how effectively the immune system performs its defensive roles. Therefore, cytokine levels indicate immunological activation rather than total immune efficacy. The decrease in these cytokines indicates that supplementation may alleviate excessive inflammation linked to placental complications, postpartum infections, or neonatal inflammatory disorders, facilitating a healthier transition for both the mare and foal. However, optimal dosing and timing require further equine-specific research.

5 Conclusions

Maternal 25(OH)D₃ supplementation in periparturient mares significantly enhanced energy metabolism and serum 25(OH)D₃ and calcium/phosphorus levels in dams and foals while increasing milk calcium, phosphorus, 25(OH)D₃, and immunoglobulin (IgA, IgG, and IgM) levels. Supplementation improved serum antioxidant capacity and anti-inflammatory responses in maternal-foal pairs, promoting dams’ and foals’ integration and healthy development. Future research should attempt to involve larger cohorts across different sites and implement standardized techniques for tracking growth measures to corroborate and broaden our results.

6 Study limitations

This study had several limitations. Primarily, the sample size was relatively small because of the challenges associated with sourcing many homogeneous experimental horses under identical management conditions. Our results may limit the statistical power and the generalizability of our findings to the broader equine population. The findings of this study, derived from a single breed and a single farm, may not be directly extrapolated to all periparturient mares under different conditions. The primary value of this work lies in providing proof-of-concept data and identifying significant response trends that warrant confirmation in larger, multi-center trials. Future studies should aim to incorporate larger cohorts across multiple sites and implement standardized protocols for monitoring growth metrics to corroborate and expand our results.

Data availability statement

The raw data supporting the conclusions of this article will be made available by the authors, without undue reservation.

Ethics statement

All experimental procedures involving animals were approved (animal protocol number: 2024019) by the Animal Welfare and Ethics Committee of Xinjiang Agricultural University, Urumqi, Xinjiang, China. The studies were conducted in accordance with the local legislation and institutional requirements. Written informed consent was obtained from the owners for the participation of their animals in this study.

Author contributions

XX: Data curation, Methodology, Conceptualization, Formal analysis, Writing – original draft, Software. MK: Conceptualization, Investigation, Validation, Software, Formal analysis, Writing – review & editing, Writing – original draft, Resources. YL: Methodology, Writing – original draft, Data curation, Resources. ML: Investigation, Writing – original draft, Data curation, Validation, Resources. JW: Resources, Writing – original draft, Formal analysis, Investigation. JH: Writing – original draft, Visualization, Funding acquisition, Data curation, Validation. FL: Funding acquisition, Writing – original draft, Investigation, Writing – review & editing, Visualization, Conceptualization, Supervision, Project administration.

Funding

The author(s) declared that financial support was received for this work and/or its publication. This research was supported by the Major Scientific and Technological Project of the Autonomous Region (Grant No. 2022A02013-2) and the Third Xinjiang Scientific Expedition Program (Grant No. 2022xjkk0404).

Acknowledgments

The authors thank every member who helped during the experiment.

Conflict of interest

The author(s) declared that this work was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

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Supplementary material

The Supplementary Material for this article can be found online at: https://www.frontiersin.org/articles/10.3389/fanim.2026.1735197/full#supplementary-material

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