Applied Animal Science 39:289–305

https://doi.org/10.15232/aas.2023-02409
© 2023, The Authors. Published by Elsevier Inc. and Fass Inc. on behalf of the American
Registry of Professional Animal Scientists. This is an open access article under the
CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

NUTRITION: Review

Review: Vitamin A supply in swine production:

Current science and practical considerations
Yauheni Shastak* and Wolf Pelletier
BASF SE, Nutrition & Health Division, Business Unit Animal Nutrition, 67063 Ludwigshafen am Rhein, Germany

ABSTRACT and health to formulate well-balanced diets and attain op-

timal performance in practical circumstances.
Purpose: This paper aims to present a comprehensive Conclusions and Applications: Retinol is crucial in
overview of vitamin A metabolism in pigs and the effects pig nutrition, and synthetic retinyl acetate supplementa-
of retinol supplementation on pig production, including tion is necessary to meet dietary requirements, as plant
immune function, reproductive performance, hematopoi- feedstuffs are deficient in vitamin A. Dietary supplemen-
esis, and antioxidant capacity. Furthermore, this review tation and injection of retinol have demonstrated posi-
highlights crucial factors that need to be considered to tive effects on swine reproduction, performance, immu-
ensure adequate vitamin A supply to pigs. nity, and health. To ensure a well-balanced pig diet, it
Sources: The primary source of information reviewed is crucial to have insights into the appropriate stages for
was independent, published scientific peer-reviewed litera- increased retinol supplementation and its interactions
ture. with other fat-soluble bioactive substances. Updates to
Synthesis: Vitamin A, also known as retinol, plays a scientific committees’ recommendations on vitamin A are
crucial role in growth, cellular metabolism, bone develop- recommended, to reflect the current state of knowledge on
ment, and reproduction. The amount of retinol supplemen- the role of vitamin A, genetics, and current pig farming
tation in commercial animal production has been a topic practices.
of contention in recent discussions. Although the physi-
ological function of vitamin A is a key factor, the cost and Key words: vitamin A, retinol, swine, supplementation,
stability of pure commercial products, along with their recommendation
stability in premix and feed formulations, are also impor-
tant considerations. Failure to provide adequate levels of INTRODUCTION
vitamin A to swine can lead to decreased performance,
increased susceptibility to diseases, and reproductive dis- When it comes to vitamins, vitamin A stands out as
orders. Determining the optimal amount of vitamin A for perhaps the most significant from a practical perspective
swine is not a simple task, as it depends on various factors (McDowell, 2000). Based on a search in the PubMed sci-
such as age, health, nutritional status, and physiological entific database (https:​/​/​pubmed​.ncbi​.nlm​.nih​.gov/​), an
state. Scientific committees such as the Gesellschaft für average of 4 articles per day have been published in the
Ernährungsphysiologie (GfE) and the National Academies last decade (from 2012 to 2022), highlighting a sustained
of Sciences, Engineering, and Medicine (NASEM) provide focus on research related to vitamin A and retinoids. This
requirement estimates (NASEM) or allowances (GfE) indicates continuing interest in the subject, even with ex-
based on the needs of healthy animals under optimal envi- isting data and knowledge already available.
ronmental conditions, serving as a foundation for practical The term “vitamin A” refers to 3 distinct chemical
considerations. However, when extrapolated to real-world compounds: retinol (the alcohol), retinal (the aldehyde),
conditions, requirement estimates and allowances derived and all-trans retinoic acid (ATRA; Olsen and Blomhoff,
on controlled research settings may have inherent limita- 2020). These compounds, collectively known as retinoids,
tions. Therefore, it becomes imperative to comprehend the get their name from their roles in the physiology of the
effects of supplemental vitamin A on swine metabolism retina (Engelking, 2014). Retinol plays an essential role
in the growth, vision, differentiation, and proliferation of
various epithelial tissues, bone formation, reproduction,
The authors have not declared any conflicts of interest. However, and embryonic development (EFSA, 2013). Throughout
both authors are employees of BASF, a manufacturer of vitamins this review, the primary focus will be on the attributed
and carotenoids, including vitamin A and β-carotene. effects of ATRA, which is the principal active form of vi-
*Corresponding author: yauheni.shastak@​basf​.com tamin A involved in swine metabolism. These effects will
290 Nutrition

Table 1. Functions of vitamin A (adapted from Combs and McClung, 2017)

Biochemical form Function

Retinol Transport
Retinyl esters Storage
Retinal Vision
All-trans retinoic acid Epithelial differentiation and growth, immune function, gene transcription, reproduction, bone
remodeling, antioxidant capacity, hematopoiesis

be thoroughly examined and discussed. Table 1 provides a This review provides an overview of the metabolism of
summary of the primary functions of vitamin A. vitamin A in pigs, as well as the influence of supplemen-
Only animal-derived feeds, such as fish liver oil, whole tal retinol on various aspects of pig production, including
milk, and fish meal, contain vitamin A. However, some reproductive performance, immune function, hematopoi-
plant pigments known as carotenoids may have vitamin esis, and antioxidant capacity. Finally, the review covers
A activity by yielding retinoids metabolically (Combs and the essential considerations for ensuring proper vitamin A
McClung, 2017). Nonetheless, only a small fraction of ca- supply in pigs.
rotenoids in nature can be transformed into vitamin A
by mammals and birds, and their utilization is reliant on METABOLISM OF VITAMIN A IN SWINE
factors such as binding form, animal species, and vitamin
A supply status (Surai, 2012). Therefore, synthetic retinol, The metabolism of vitamin A in swine is a complex and
most commonly in the form of retinyl acetate, is typically multistep process that involves several enzymes and path-
used in animal nutrition to fulfill dietary demand for vi- ways. Ingested primarily as retinyl esters such as retinyl
tamin A. acetate, vitamin A is hydrolyzed by pancreatic lipases and
Failure to provide adequate vitamin A in animal nutri- esterases in the small intestine to release retinol, which
tion may result in growth and bone formation disorders, is then incorporated into mixed micelles with other lip-
skin and mucosal pathological changes, and impaired visu- ids and bile salts to facilitate its efficient absorption by
al ability, including night blindness. Moreover, deficiency enterocytes (Hynd, 2019). Once absorbed, retinol is re-
in vitamin A may increase susceptibility to diseases and esterified and integrated into chylomicrons that trans-
cause reproductive disorders. port vitamin A to the liver via the lymphatic system and
The optimal amount of retinol and its metabolites re- bloodstream (Harrison, 2012). In the liver, vitamin A is
quired to promote performance and overall health may stored as retinyl esters in hepatic stellate cells, which can
vary depending on various factors. When formulating diets, be mobilized when needed (Carazo et al., 2021). Upon
natural levels of vitamin A and carotenoids in feedstuffs mobilization, retinyl esters are hydrolyzed to retinol and
are typically not taken into consideration, and instead, the bound to retinol-binding protein for transport to target
total retinol requirement is met through supplementation tissues (Steinhoff et al., 2022). Retinol-binding protein is
(Darroch, 2000). In swine diets, vitamin A is often supple- a critical regulator of vitamin A homeostasis, as it ensures
mented at higher doses than recommended by GfE (2006) proper distribution and delivery of vitamin A to target
and NASEM (2012), as it may provide additional benefits tissues while preventing toxicity.
such as enhancing epithelium development, improving im- Once retinol reaches the target tissue, it can be metabo-
munity and overall health in growing-finishing pigs, and lized into active forms, such as ATRA, which is involved
optimizing sow reproductive performance (Lindemann et in regulating gene expression and cellular differentiation
al., 2008; Olivares et al., 2009). (Zasada and Budzisz, 2019). The conversion of retinol to
It is worth noting that most nutritionists consider the ATRA is mediated by several enzymes, including alcohol
NASEM vitamin requirements to be close to the minimum dehydrogenases and retinal dehydrogenases (Bchini et al.,
requirements necessary to prevent clinical deficiency (Mc- 2013). These enzymes are essential for proper vitamin A
Dowell, 1989; Kırkpınar and Açıkgöz, 2018). Moreover, it metabolism and play critical roles in determining the bio-
is important to keep in mind that retinyl acetate may in- availability and physiological effects of vitamin A in swine.
cur significant losses during storage and processing, which According to Green and Fascetti (2016) and Hynd
are not taken into account in scientific vitamin standards (2019), retinaldehyde can be synthesized from carotenoid
(Hirai et al., 2023). McDowell (2006) suggests that the precursors, which are absorbed through passive diffusion
optimum vitamin supplementation level is the amount and subsequently cleaved by dioxygenases in the intestinal
that results in the best growth rate, feed utilization, and mucosal cells, indicating the potential for the formation
health, including immune competency, while also provid- of vitamin A. Retinaldehyde is converted to ATRA via a
ing sufficient body reserves. 2-step oxidation process (Belyaeva et al., 2019). The first
 Shastak and Pelletier: Vitamin A supply in swine production 291

step is catalyzed by retinaldehyde dehydrogenase, which Vitamin A as retinal plays a critical role in the visual
adds a carboxyl group to the aldehyde group. The second system of animals, as it is a key component of the visual
step is carried out by a group of enzymes called aldehyde pigment rhodopsin. Rhodopsin is present in the retinal
dehydrogenases, resulting in the formation of retinoic acid. rods and is essential for vision under low-light conditions,
The conversion of carotenoids to vitamin A in pigs is triggering a signaling cascade that ultimately results in vi-
inefficient and highly variable. Swine show limited abil- sual perception in swine (Zhou et al., 2011). This pigment
ity to convert carotenoids to vitamin A, probably due to is composed of 2 elements: opsin, a protein, and 11-cis-
the low activity of the enzyme responsible for this con- retinal, the chromophore derived from retinol (Filipek et
version, β-carotene 15,15′-dioxygenase. Conversion rates al., 2003). Night blindness and complete blindness in ani-
of β-carotene to vitamin A in pigs have been reported mals can occur due to a deficiency of vitamin A (Debelo
to range from 7:1 to 40:1 (Ullrey et al., 1965; NASEM, et al., 2017).
1998; Schweigert et al., 2001), with an average conver- The subsequent sections of this scientific review paper
sion rate that is considered the lowest among domestic will provide a comprehensive analysis of the effects of vi-
animals. Thus, typically, 22 μg of β-carotene is required tamin A supplementation on various physiological func-
to form 1 μg of retinol or 3.33 IU of vitamin A in swine. tions, including reproductive performance, immune func-
Although β-carotene has the potential to serve as a source tion, hematopoiesis, and oxidative capacity. It is worth
of vitamin A in swine, its contribution is hindered by fac- mentioning that, in this manuscript, the term “vitamin A
tors such as the highly variable content of β-carotene in deficiency” specifically pertains to diets or animals that
feedstuffs, limited and unpredictable conversion rates to are not supplemented with any source of retinol or that
retinaldehyde, low stability, and the effects of animal age, are provided with levels below the requirement estimates
genetics, stress, physiological state, and health-related fac- of NASEM (1988, 1998, 2012).
tors. Consequently, accurately estimating the amount of
vitamin A that can be obtained from β-carotene in pig EFFECTS ON REPRODUCTIVE
diets is challenging. For a more detailed investigation of PERFORMANCE OF SOWS
this topic, readers are referred to the excellent work of
Green and Fascetti (2016), which provides a comprehen- One of the main obstacles to profitability in the pork
sive analysis of the same phenomenon. industry is the reproductive efficiency of breeding sows.
The biologically active form of vitamin A, ATRA modu- Retinol is a crucial vitamin that plays a vital role in main-
lates cellular metabolism in swine by binding to specific taining the reproductive system and promoting fetal de-
nuclear receptors, namely retinoic acid receptors and reti- velopment (Thompson et al., 1964). Through specific in-
noid X receptors, which are widely expressed in various teractions with cellular receptors, ATRA, the active form
tissues, including liver, kidney, intestinal, and adipose of vitamin A, plays a crucial role in various reproductive
tissues and immune cells (Brandebourg and Hu, 2005). processes. In the ovaries, it influences granulosa cell prolif-
Upon binding, the retinoic acid-receptor complex interacts eration, differentiation, and steroidogenesis, contributing
with specific DNA sequences, leading to the regulation to follicular development (Demczuk et al., 2016; Jiang et
of gene expression in diverse biological processes, such as al., 2018). Further, ATRA stimulates the expression of
vision, immunity, and cell differentiation (Rastinejad et genes involved in reproduction and enhances the produc-
al., 2000). For instance, ATRA plays a vital role in the tion of estrogen, which is essential for estrus synchroni-
differentiation of intestinal epithelial cells, the maturation zation and oocyte maturation (Munetsuna et al., 2009;
of immune cells, and the regulation of adipose tissue me- Damdimopoulou et al., 2019). Additionally, ATRA modu-
tabolism. A recent investigation conducted by Wang et al. lates uterine receptivity and embryo implantation, pro-
(2020) demonstrated that vitamin A has the potential to moting the secretion of uterine factors that aid in embryo
modulate growth performance and intestinal function in attachment and placental development (Han et al., 2010;
weaned piglets by regulating intestinal stem cells located Yin et al., 2021). Moreover, ATRA plays a crucial role in
in the jejunum. male reproduction, influencing spermatogenesis, testicular
Vitamin A, specifically ATRA, plays a critical role in development, and sperm production and motility (Crespo
bone growth and development in addition to its cellu- et al., 2019). These cellular effects of vitamin A underline
lar differentiation functions. Retinol deficiency has been its significance in optimizing fertility, litter size, and over-
found to impair bone growth and increase the risk of bone all reproductive performance in swine.
malformations in pigs (Palludan, 1961). The production Coffey and Britt (1993) were pioneers in demonstrating
and activation of osteoblasts, responsible for building that litter size could be increased through the injection
new bone tissue, can be partially attributed to the role supplementation of retinyl palmitate to sows who were
of vitamin A (Chiba et al., 1996). Additionally, retinol already receiving adequate dietary levels of vitamin A in
is involved in the regulation of bone resorption, which is the form of retinyl acetate (11,000 IU/kg of feed). The
the natural process of breaking down old bone tissue and treatments consisted of intramuscular administration of
replacing it with new bone tissue (Yee et al., 2021). 50,000 IU of vitamin A or vehicle on the day of wean-
292 Nutrition

ing, on the day of mating, and on d 7 after mating. The (1993) and Silveira et al. (1997). In their research, Linde-
study found that injection with vitamin A resulted in an mann et al. conducted injections of either 250,000 IU or
increase in the number of piglets born alive and total litter 500,000 IU of vitamin A into sows at weaning and breed-
weight. These findings suggest that injection supplemen- ing, while these sows were already receiving a diet with
tation with vitamin A may have positive effects on litter sufficient levels of vitamin A (11,000 IU/kg of feed). The
size and weight, even in sows receiving adequate levels of study involved a large sample of 443 litters from 182 sows
supplemental vitamin A in their diets. The authors con- across 5 experimental stations. Results showed a con-
cluded that the observed favorable responses in sows, fol- sistent increase in litter size for both total piglets born
lowing treatment with vitamin A, suggest an essential role and piglets weaned per litter, with a linear correlation
for this micronutrient in early embryonic development. to the amount of vitamin A palmitate that was injected.
In a series of studies conducted by Robertson (1997), the The findings of this comprehensive study emphasize the
effectiveness of retinyl palmitate and retinyl propionate critical role of maternal vitamin A supplies in the devel-
in improving reproductive parameters in swine, such as opment and performance of offspring. Lindemann et al.
follicle development, was compared. The studies involved (2008) have suggested that vitamin A enhances litter size
administering a 1,000,000-IU i.m. injection of retinyl pal- by positively affecting various reproductive processes such
mitate on d 16 of the estrous cycle and a 500,000-IU i.m. as oocyte maturation, ovulation, fertilization, early em-
injection of retinyl propionate on the day before weaning bryonic survival, and development.
(group 1), at day of mating (group 2), or 6 d after mating Whaley et al. (2000) administered 1,000,000 IU of reti-
(group 3). The results showed that retinyl palmitate had nyl palmitate on d 15 after second estrus to gilts fed high-
a positive effect on follicle development, possibly through energy diets, and found that it altered the development of
the steroidogenesis pathway. However, it did not appear oocytes and embryos. Vitamin A increased the mean stage
to affect the expression of IGF-1 in embryonic tissue. On of development, decreased the variation in stage, and al-
the other hand, the injection of retinyl propionate did not tered the follicular hormonal environment. The authors
have any noticeable effects on ovulation rate, IGF-1 lev- concluded that vitamin A may influence embryonic devel-
els, or the formation of corpora lutea. Interestingly, the opment by advancing the resumption of meiosis.
author suggested that retinyl palmitate was more effective Tonn et al. (1994) conducted a study to evaluate the
than retinyl propionate in enhancing reproductive traits in effects of administering vitamin A (1,000,000 IU) or go-
swine. However, this conclusion raises questions because nadotropin injection to sows at weaning. The study as-
retinyl propionate was administered at a dosage 2 times sessed the ovulation rate and early embryonic survival
lower than retinyl palmitate (500,000 IU vs. 1,000,000 IU), and development 11.5 d after the onset of estrus. The
and there was also difference in the timing of the injec- results showed that both vitamin A and gonadotropin sig-
tions. Therefore, further studies are necessary to deter- nificantly increased the number of embryos recovered. In-
mine whether there are indeed differences in the biological terestingly, the 2 treatments appeared to enhance embryo
value of different intramuscularly injected retinyl esters. recovery through different mechanisms. Specifically, sows
Furthermore, it is worth noting that Robertson (1997) treated with gonadotropin injections displayed a greater
used only a single dose of vitamin A in their study. number of ovulated eggs, whereas those administered with
Silveira et al. (1997) conducted a field trial with 1,030 vitamin A demonstrated a tendency toward elevated em-
sows to specifically examine the effects of vitamin A injec- bryo survival rates.
tion (450,000 IU of retinyl palmitate) at 3 time points: However, it is worth noting that certain studies have re-
at weaning, during mating, and 7 d after mating. The ported no significant effects of vitamin A on the reproduc-
objective was to investigate the influence of this vitamin tive performance of sows (Tokach et al., 1994; Pusateri et
A treatment on the subsequent litter size of multiparous al., 1999). In the study by Tokach et al. (1994), sows were
sows at weaning or breeding. The study found that the administered 1,000,000 IU of vitamin A (retinyl propio-
administration of retinyl palmitate had a positive effect on nate; i.m.) at weaning, whereas Pusateri et al. (1999) in-
the reproductive performance of sows, regardless of their jected sows with 1,000,000 IU of vitamin A (specific form
productive phase. Specifically, it led to an increase in the not mentioned) at various time points: at weaning or on d
number of total and live-born piglets, as well as an in- 0, 2, 6, 10, 13, 19, 30, 70, or 110 after breeding. Both To-
crease in litter weight at birth. In light of these findings, kach et al. (1994) and Pusateri et al. (1999) acknowledged
the authors recommended the administration of 500,000 that the lack of observed effects in their studies could be
IU of vitamin A palmitate to sows at weaning or to gilts 5 attributed to the need for prolonged exposure to vita-
d before the expected heat period. Collectively, the study min A, through either multiple injections or the use of a
highlights the potential benefits of vitamin A administra- sustained-release form. Additionally, Tokach et al. (1994)
tion in improving the reproductive outcomes of sows, and noted a significant number of piglets born alive to the con-
offers practical recommendations for its use in pig farm- trol sows (without vitamin A injection), which may have
ing. affected the study results. Furthermore, incorrect timing
In support of this, Lindemann et al. (2008) conducted of the injections was also mentioned as a potential factor.
a study that confirmed the findings of Coffey and Britt Lindemann et al. (2008) also emphasized that a single
 Shastak and Pelletier: Vitamin A supply in swine production 293

Table 2. Overview of the influence of vitamin A injection supplementation in sows on litter size of live-born piglets

Litter size of piglets born

Dietary vitamin alive—control group Litter size of piglets
A level, (no vitamin A injection from sows injection
Study IU/kg of feed supplementation in sows) supplemented with vitamin A
Coffey and Britt (1993) 11,000 10.0 10.6*
Robertson (1997) 15,000 10.6 10.41
Silveira et al. (1997) “Commercial level” 10.3 10.7*
Lindemann et al. (2008) 11,000 9.7 11.1,2* 12.23*
Tokach et al. (1994) 10,000 10.4 10.51
Pusateri et al. (1999) Unknown 10.1 10.11
Brief and Chew (1985) None4 7.9 9.5*
1
Single injection of vitamin A.
2
Intramuscular injection with 250,000 IU of vitamin A.
3
Intramuscular injection with 500,000 IU of vitamin A; 11,000 IU per kg of diet.
4
To conduct comparative analysis between dietary supplementation and injection supplementation at an equivalent level.
*Significantly different from the control group (P < 0.05).

injection of vitamin A, as seen in the studies by Tokach et sage of retinyl esters through the digestive system. This
al. (1994) and Pusateri et al. (1999), or short-term retinol process includes degradation of the esters by digestive
supplementation, can lead to inconclusive effects. Thus, to enzymes, their absorption across the intestinal wall, and
obtain definitive results, longer exposure to vitamin A is their subsequent metabolism in the liver before reaching
necessary. The effects of vitamin A injection supplementa- systemic circulation. This multistep process highlights the
tion in sows on litter size of live-born piglets are presented challenges faced by retinyl esters supplied orally, poten-
in Table 2, providing an overview of the findings. tially resulting in lower biological value compared with
When examining the influence of vitamin A on reproduc- injection methods. Generally, i.m. injections have a faster
tive performance, it is crucial to take into account poten- onset of action compared with s.c. injections, due to the
tial variations in absorption and bioavailability depending greater vascularity and blood flow in muscle tissue (Pola-
on the route of administration, which can subsequently nia Gutierrez and Munakomi, 2023). However, s.c. injec-
affect an animal’s overall vitamin A status. tions may have a more prolonged duration of action com-
Brief and Chew (1985) conducted a study with gilts that pared with i.m. injections because the active substance
were given either dietary supplementation of vitamin A or is absorbed and released more slowly from subcutaneous
injected with vitamin A propionate for 4 weeks before far- tissue (Rahnfeld and Luciani, 2020). It is important to
rowing. Injected gilts had lower embryonic mortality and note that different retinyl esters may exhibit distinct ab-
produced more and heavier piglets per litter than those sorption characteristics based on their specific properties
that were fed or deficient in vitamin A. The study suggests (e.g., molecular weight, stability) and the site of injection.
that injection supplementation of vitamin A is more effec- Finally, administration of normal or oxidized non-provi-
tive than dietary supplementation in improving reproduc- tamin A β-carotene through dietary or injection supple-
tive performance in gilts. mentation has resulted in more inconsistent effects on re-
The route of vitamin A administration clearly indicates productive traits in gilts and sows compared with vitamin
variations in the biological values of different adminis- A, as demonstrated by various studies (Brief and Chew,
tration types in animals. For instance, direct injection of 1985; Coffey and Britt, 1993; Sternder et al., 1999; Kosto-
retinyl acetate into the bloodstream yields a reference bio- glou et al., 2000; Chen et al., 2021; Elefson et al., 2023).
availability value of 100%, whereas other methods such as Although β-carotene is believed to have beneficial effects
i.m. or s.c. injection and oral supplementation (via feed or on reproduction, this variability could be attributed to
water) have lower bioavailability values. Direct injection several factors, such as the varying levels of conversion of
of retinyl esters into the bloodstream bypasses the barri- normal β-carotene to vitamin A, its independent effect on
ers and tissues encountered with other routes, resulting in reproduction, variable bioavailability, route of administra-
faster and more complete utilization. When retinyl esters tion, and dosage levels.
are administered through s.c. or i.m. injections, they need In conclusion, vitamin A injection supplementation has
to diffuse across cellular barriers and blood vessels before been found to have positive effects on the reproductive
entering systemic circulation. Oral administration of vita- performance of breeding sows, such as increases in litter
min A, whether through feed or water, involves the pas- size and weight, and enhancement of various reproductive
294 Nutrition

processes. The results of these studies suggest that vita-

min A administration during crucial developmental stages
of the fetus could potentially improve sow reproductive
performance. However, it is important to note that some
studies have reported no effect of vitamin A on reproduc-
tive performance parameters of sows, which highlights the
need for further research in this area. Overall, the use
of vitamin A supplementation could be a valuable tool
for pig farmers to improve the reproductive efficiency of
breeding sows.

EFFECTS ON IMMUNE SYSTEM Figure 1. Treatment of pigs with vitamin A increased eosinophils
in lung tissue after inoculation with the large roundworm
Vitamin A is a micronutrient that has been widely stud- (adapted from Dawson et al., 2009). Treatments annotated with
ied for its role in immune function (Villamor and Fawzi, unique letters (a–c) are statistically different (P < 0.05).
2005). In fact, as early as 1928, Green and Mellanby dis-
covered the crucial role that vitamin A plays in immune supplemented with vitamin A. The study found that the
modulation, naming it “the anti-infective vitamin” (Green deficient pigs generated less than one-tenth of the amount
and Mellanby, 1928, page 691). It is now well-established of antibodies compared with the control group receiving
that vitamin A not only helps to maintain the integrity of vitamin A supplements. These findings underscore the im-
mucosal epithelial barriers but also has direct effects on portant role of vitamin A in supporting robust antibody
the function of various immune cells (Engedal, 2011). production.
At the cellular level, ATRA has been shown to regulate Chew (1987) stated that retinol supplements can poten-
immune cell differentiation, proliferation, and function tially improve resistance against both gram-negative and
(Ross, 2012; Brown and Noelle, 2015), exerting its immu- gram-positive bacteria, and also mitigate parasitic infec-
nomodulatory effects through interactions with retinoic tions. Wagner and Polley (1997) have pointed out that
acid receptors (RAR) and retinoid X receptors (RXR) over 50% of fattening swine worldwide are infected by the
expressed on various immune cell populations (Zisakis et large roundworm of pig. Studies have shown that vitamin
al., 2007). Further demonstrated effects of ATRA include A can enhance the localized immune response in mam-
the ability to enhance natural killer (NK) cell production mals infected with parasitic nematodes (Long et al., 2006;
and activity, promoting cytotoxicity against infected or Payne et al., 2007; Dawson et al., 2009). Dawson et al.
transformed cells (Grudzien and Rapak, 2018). Further- (2009) have shown that vitamin A supplementation, in the
more, ATRA influences the development and differentia- form of retinoic acid, can increase the expression of mark-
tion of T cells, regulating cytokine production and pro- ers for T lymphocytes (including Th1, Th2, and regula-
moting the generation of regulatory T cells (Tregs) crucial tory T cells) in the liver and lungs, as well as increase
for immune tolerance (Bono et al., 2016). It also affects eosinophilia in the lungs of large roundworm-infected pigs
the function of antigen-presenting cells, such as dendritic (Figure 1). These findings suggest that vitamin A sup-
cells, enhancing their antigen-capturing and presentation plementation may potentially stimulate a robust immune
abilities to T cells (Geissmann et al., 2003). Additionally, response that could be sufficient to control parasites and
ATRA contributes to maintaining mucosal immune ho- limit inflammation, depending on the specific situation.
meostasis by promoting the production of secretory immu- Even minor deficiencies in vitamin A can have significant
noglobulin A in the gut-associated lymphoid tissue (Mora effects on both innate and adaptive immune responses, ul-
and von Andrian, 2009). Moreover, the retinol metabolite timately increasing susceptibility to infections (Vlasova et
14-hydroxy-4,14-retro-retinol (14-HRR) serves as a vital al., 2013). For example, studies have shown that piglets
cofactor for B lymphocyte growth and T lymphocyte acti- deficient in vitamin A and infected with rotavirus experi-
vation (Buck et al., 1993). Notably, the ability to convert enced prolonged fecal shedding of the virus, higher levels
all-trans retinol to 14-HRR is present in almost all cells of virus in their feces, and more severe diarrhea following
across species, indicating its conservation from insects to infection compared with piglets who were supplemented
mammals (Blomhoff and Blomhoff, 2006). Vitamin A also with vitamin A. These findings highlight the crucial role
plays a crucial role in the synthesis of antiviral substances of vitamin A in supporting immune function and protect-
such as lysozyme (West et al., 1991; Zhou et al., 2021). ing against infectious disease. In general, earlier or longer
Thus, compelling evidence underscores vitamin A’s essen- vitamin A supplementation of both mothers and their off-
tiality for optimal immune function, serving a critical role spring is needed to improve intestinal health and normal-
in defending the body against infections and diseases. ize immune response in swine (Vlasova et al., 2013).
Early research conducted by Harmon et al. (1963) re- Chattha et al. (2013) found that vitamin A deficiency
vealed a significant disparity in antibody production be- in pigs negatively affects adaptive B and T lymphocyte
tween pigs deficient in vitamin A and control animals responses and causes imbalances in innate and adaptive
 Shastak and Pelletier: Vitamin A supply in swine production 295

cell distribution. Vitamin A-deficient pigs showed higher injections are commonly administered to piglets to treat
CD8 T cell frequencies and lower CD4 T cell frequencies this disorder. However, it is worth noting that anemia can
in various tissues after exposure to a virulent rotavirus. also arise from a deficiency in vitamin A.
The researchers observed increased plasmacytoid and con- Research findings have demonstrated the significant in-
ventional dendritic cells before the challenge, indicating an volvement of retinoids and ATRA in the differentiation
overall disruption of immune responses. Compared with and maturation of hematopoietic stem cells (HSC) within
vitamin A-supplemented pigs, unvaccinated vitamin A- the bone marrow (Purton, 2007). The HSC effectively
deficient piglets had lower hepatic vitamin A levels, more maintain hematopoietic fitness by carefully balancing
severe and prolonged diarrhea, and greater rotavirus shed- self-renewal and differentiation throughout an organism’s
ding post-challenge. Vitamin A-deficient pigs also had lifespan. The study by Menéndez-Gutiérrez et al. (2023)
higher levels of innate cytokines (IFNα) and CD8 T cells, highlights the critical role of RXRα and RXRβ in sustain-
along with elevated proinflammatory cytokines (IL12) and ing HSC quiescence, survival, and chromatin structure.
decreased anti-inflammatory cytokines (IL10). Orally vac- Deficiencies in these receptors lead to altered gene expres-
cinated vitamin A-deficient pigs had lower serum IgA rota- sion patterns resembling aged HSC, upregulation of the
virus antibody titers, higher diarrhea severity scores, and MYC (myelocytomatosis oncogene) pathway, and changes
significantly fewer intestinal IgA antibody-secreting cells in RNA splicing. Furthermore, ATRA has been shown to
post-challenge than vaccinated vitamin A-supplemented promote the commitment of multipotent progenitors to-
pigs, indicating lower anamnestic responses. ward the myeloid and lymphoid lineages, ultimately con-
Recently, Chepngeno et al. (2022) reported that admin- tributing to the production of erythrocytes, leukocytes,
istering rotavirus (maternal immunization) to pregnant and platelets (Rönn et al., 2015).
sows resulted in increased lactogenic immunity and pas- According to Combs and McClung (2017), the hema-
sive protection in piglets. The researchers also observed tological response to vitamin A deficiency has 2 phases:
that supplementing vitamin A during gestation and lacta- first, a decline in hemoglobin and erythrocyte count due
tion improved sow immune responses and passive protec- to impaired hemoglobin synthesis, followed by an increase
tion in offspring. These results provide insight into the role in both variables due to hemoconcentration caused by
of retinol in infection and vaccination, the effects of ma- dehydration and diarrhea. In human studies, vitamin A
ternal vitamin A deficiency and immunization on neonatal supplementation during pregnancy has been found to help
passive immune protection, and the potential to enhance control nutritional anemia, and combined therapy with
swine rotavirus vaccination programs to improve control retinol and iron has produced synergistic effects (Suharno
of neonatal rotavirus infection. et al., 1992, 1993; Jiang et al., 2009). The metabolic im-
The effects of vitamin A supplementation on preg- portance of vitamin A in hematopoiesis seems to involve
nant swine infected with porcine epidemic diarrhea virus the mobilization and transport of iron from body reserves,
(PEDV) were investigated by Langel et al. (2019). Their as well as an increase in nonheme-iron bioavailability
study revealed that vitamin A supplementation during the (Combs and McClung, 2017).
third trimester of pregnancy led to improvements in gut According to an experimental study conducted on neo-
homeostasis, immune regulation, and anti-PEDV IgA im- natal piglets, the combination of iron and vitamin A is
munity in the blood, milk, and ileum. These positive out- more effective in improving iron status compared with us-
comes resulted in increased survival rates among PEDV- ing iron alone (Jiang et al., 2009). The study found that
challenged litters. The findings of this study suggest that administering intramuscular injections of iron in combina-
vitamin A supplementation may have the potential to tion with oral vitamin A (2,000 IU vitamin A as vitamin
enhance intestinal immunity during pregnancy and lacta- A palmitate at d 2 after birth) was more effective than
tion, and could be used to inform maternal vaccination using iron alone. The results showed that on d 7 and 14,
and retinol supplementation strategies for enteric viral dis- plasma iron concentrations were significantly higher in
eases in both humans and animals. piglets treated with both iron and vitamin A compared
Thus, vitamin A is crucial for immune modulation, with those treated with iron alone. Levels of hemoglobin
maintaining mucosal barriers, enhancing mucosal homing, and hematocrits were used to evaluate the body’s biologi-
generating regulatory T cells, and increasing IgA secre- cal response to iron, and the study found that on d 7, 14,
tion. Deficiencies can lead to increased susceptibility to and 21 after birth, hemoglobin concentrations were 11.2%,
infections and diseases, whereas supplementation can im- 10.6%, and 10.7% greater, respectively, in piglets treated
prove immune response, limit inflammation, and enhance with both iron and vitamin A than in piglets treated with
vaccine efficacy. iron alone. Based on these findings, the authors concluded
that vitamin A supplementation in piglets may enhance
EFFECTS ON HEMATOPOIESIS the role of iron in preventing piglet anemia.
Swine have been used as research models in various ar-
Neonatal iron deficiency anemia is a common condition eas of medical and biological research due to their ana-
in swine, the only mammalian species known to suffer from tomical, physiological, and genetic similarities to humans.
this problem (Szudzik et al., 2018). Iron supplements or Conversely, studies conducted in humans can also provide
296 Nutrition

valuable information for pig research. These reciprocal ways, downregulating NADPH oxidase, a key enzyme
benefits between human and pig research highlight the involved in reactive oxygen species production (Azzam
mutual value in studying both species. et al., 2022). Tourniaire et al. (2015) demonstrated that
Saraiva et al. (2014) studied the relationship between ATRA upregulates genes involved in mitochondrial bio-
serum retinol levels and anemia and iron deficiency in chil- genesis and antioxidant defense, leading to an increase
dren aged 1 to 5 years. They collected dietary intake data in oxidative phosphorylation capacity and mitochondrial
and biochemical parameters including ferritin, hemoglo- content. Finally, ATRA has been shown to induce au-
bin, and retinol serum. The study found that children with tophagy, a cellular process that enables cells to survive in
retinol deficiency had a significantly greater prevalence conditions of nutrient depletion or oxidative stress (Choi
of anemia and iron deficiency. The association remained et al., 2013; Fang et al., 2020). According to Rajawat et
significant even after adjusting for socioeconomic and de- al. (2010), ATRA promotes the maturation of autophago-
mographic conditions, dietary intake, and anthropometric somes through a pathway that does not involve the classic
variables. Furthermore, positive correlations were found nuclear retinoid receptors. Retinoic acid causes a redistri-
between serum retinol levels and both serum ferritin and bution of a receptor called cation-independent mannose-
hemoglobin levels. The study highlights the importance of 6-phosphate receptor from the trans-Golgi region to the
adequate vitamin A intake in preventing anemia and iron maturing autophagosomal structures, which induces their
deficiency in young children. acidification. Autophagy has been implicated in the regu-
In conclusion, although iron supplements are commonly lation of redox imbalance and the maintenance of cellu-
used to treat neonatal iron deficiency anemia in pigs, it is lar homeostasis. In a recent study by Saurav and Manna
important to consider the role of vitamin A in hematopoi- (2022), it was demonstrated that ATRA induces the ex-
esis. Limited studies have shown that combined therapy pression of profilin, an actin regulator. Furthermore, their
with iron and vitamin A may produce synergistic effects research revealed that profilin promotes autophagy by
in preventing piglet anemia. Similarly, studies in human inhibiting mTOR, upregulating the autophagy-activating
populations have shown a significant association between kinase ULK1 and stabilizing and activating AMPK (5′
retinol deficiency and anemia and iron deficiency in young adenosine monophosphate-activated protein kinase). In
children. Although further research is needed to under- conclusion, the diverse effects of ATRA on cellular anti-
stand the precise mechanisms underlying the relationship oxidant defense mechanisms, oxidative stress pathways,
between vitamin A and iron status, these findings suggest mitochondrial function, and autophagy highlight its po-
that vitamin A supplementation may enhance the effec- tential as a multifaceted regulator of redox balance and
tiveness of iron therapy in preventing anemia. cellular homeostasis.
In a study by Hu et al. (2020) involving weaned pig-
EFFECTS ON ANTIOXIDANT CAPACITY lets, the researchers examined the effects of gelatin- and
starch-encapsulated vitamin A on growth performance
The efficacy of vitamin A as an antioxidant in animal and antioxidant capacity. The results showed noteworthy
cells has been a subject of debate for some time. Combs differences between the 2 forms of vitamin A. On d 21
and McClung (2017) suggested that retinol was unlikely of the experiment, the piglets supplemented with starch-
to have a direct effect on body systems protecting against encapsulated vitamin A exhibited a significant increase
the adverse effects of pro-oxidants, as retinol and reti- in glutathione peroxidase levels compared with those re-
nal were not able to quench singlet oxygen and had only ceiving gelatin-encapsulated vitamin A. However, on d 42,
weak capability to scavenge free radicals. However, recent the response was reversed, with the group given gelatin-
studies have revealed that the antioxidant defense system encapsulated vitamin A showing significant improvement
mediated by retinol is much more intricate, involving the in glutathione peroxidase levels compared with the control
upregulation of genes related to antioxidants, heightened group, whereas no significant change was observed in the
activity of antioxidant enzymes, and improved mitochon- group receiving starch-encapsulated vitamin A.
drial function (Gad et al., 2018; Hao et al., 2021). The The study by Hu et al. (2020) sheds light on the distinct
primary mediator of vitamin A’s antioxidant effects is its effects of different sources of vitamin A in piglets’ perfor-
metabolite, ATRA. mance. Specifically, the piglets responded more favorably
All-trans retinoic acid, ATRA, has been reported to to starch-encapsulated vitamin A. This finding has practi-
enhance cellular antioxidant defense mechanisms by up- cal implications, as it suggests that the biological value of
regulating the expression of glutathione-related genes, vitamin A can vary depending on the formulation of reti-
including glutathione peroxidase and glutathione reduc- nyl acetate. Nevertheless, the results reported by Hu et al.
tase (El Haddad et al., 2012; Brigelius-Flohé and Flohé, (2020) indicate that dietary supplementation of vitamin
2020). Moreover, ATRA has been found to increase su- A has the potential to enhance antioxidant capacity and
peroxide dismutase activity, a critical antioxidant enzyme improve growth performance.
that neutralizes superoxide radicals (Rao et al., 2010). In A similar outcome was reported by Zhou et al. (2021),
addition to these effects, ATRA modulates the expres- where 2 different sources of vitamin A led to a significant
sion of several genes involved in oxidative stress path- improvement in the antioxidant capacity of piglets during
 Shastak and Pelletier: Vitamin A supply in swine production 297

Table 3. Effects of 2 different sources of vitamin A on the growth performance of weaned

piglets fed experimental diets for 35 d (Zhou et al., 2021)1

Parameter NC EQ Vit A SA Vit A SEM P-value

Average initial weight, kg, d 1 8.32 8.34 8.36 0.56 0.965
Average final weight, kg, d 35 27.23a 27.94b 28.11b 0.16 0.035
ADFI, g/d 825 830 832 2.56 0.533
ADG, g/d 540b 560a 564a 4.17 0.021
F/G 1.53 1.48 1.48 0.12 0.100

Mean values within a column with no common superscript differ significantly (P < 0.05).
a,b

1
NC = basal diet without supplemental vitamin A; EQ Vit A = basal diet supplemented with
12,000 IU/kg vitamin A stabilized with ethoxyquin; SA Vit A = basal diet supplemented with
12,000 IU/kg vitamin A stabilized with sodium ascorbate.

the 1- to 36-d postweaning period. As a result, perfor- OPTIMIZING VITAMIN A SUPPLY IN SWINE
mance also improved (Tables 3 and 4).
Over the last 2 decades, the findings of several studies Overall, the aforementioned effects highlight the poten-
have shown that the use of supplemental retinol can have tial benefits of vitamin A supplementation or injection for
a positive influence in mitigating the negative effects of improving swine performance and well-being. Providing
oxidative stress in domestic fowl, beef, and dairy cattle. commercial pig herds with the right amount of vitamin A
The studies conducted by Kucuk et al. (2003), Ma et al. is essential for achieving the best growth rate, feed utiliza-
(2005), Jin et al. (2014), and Liang et al. (2019) have con- tion, and health.
tributed to the growing body of evidence supporting the In times of high vitamin A prices, some nutritionists
effectiveness of supplemental retinol in mitigating oxida- may reduce retinyl acetate levels in feed to match the
tive stress. recommendations of scientific committees such as NASEM
In contrast, some studies have recently suggested that (2012) or GfE (2006), believing that these recommenda-
high levels of vitamin A supplementation may have a par- tions are sufficient for practical conditions. However, it
adoxical effect on rats submitted to aerobic exercise, lead- is important to note that these guidelines are based on
ing to an increase in oxidative stress and negative health ideal environmental conditions and may not account for
outcomes (Gasparotto et al., 2015; Petiz et al., 2017). individual variation in vitamin A requirements, which can
Thus, it is important to note that further research is be influenced by factors such as genetics, age, sex, body
still needed to fully understand the mechanisms behind weight, and contextual parameters. Therefore, require-
the positive effects of retinol supplementation on redox ment estimates and allowances from controlled research
imbalance and to determine the optimal dosage and ap- settings may have limitations when applied to real-world
plication methods for different animal species and produc- conditions (e.g., stocking density, disease pressure, genet-
tion systems. Nonetheless, the current evidence suggests ics), requiring careful consideration and possible adjust-
that retinol supplementation could be a valuable tool for ments. To avoid potential deficiencies, it is recommended
reducing oxidative stress in a range of contexts. to consult with a qualified nutritionist or veterinarian

Table 4. Effects of 2 different sources of vitamin A on serum biochemical parameters of weaned

piglets on d 36 of the study (Zhou et al., 2021)1

Parameter NC EQ Vit A SA Vit A SEM P-value

Serum retinol, μg/L 136b 208a 219a 12.44 0.001
Total antioxidant capacity, U/mL 1.60b 1.50b 3.36a 0.29 0.001
Glutathione peroxidase, U/mL 696b 837a 778a 21.29 0.028
Superoxide dismutase, U/mL 52.91 54.97 64.76 2.59 0.132
Malondialdehyde, nmol/mL 2.94a 1.75b 1.60b 0.22 0.008

Mean values within a column with no common superscript differ significantly (P < 0.05).
a,b

1
NC = basal diet without supplemental vitamin A; EQ Vit A = basal diet supplemented with
12,000 IU/kg vitamin A stabilized with ethoxyquin; SA Vit A = basal diet supplemented with
12,000 IU/kg vitamin A stabilized with sodium ascorbate.
298 Nutrition

Table 5. National Academies of Sciences, Engineering, and Medicine (NASEM) vitamin A requirement estimates and
Gesellschaft für Ernährungsphysiologie (GfE) vitamin A allowances

Vitamin A requirement, IU/kg of feed

Committee Piglets Finishing gilts and barrows Gestating gilts and sows Lactating gilts and sows
NASEM, 19591 1,323–1,764 882 2,939 2,939
NASEM, 19641 1,764–2,205 1,323 3,307 3,307
NASEM, 19681 1,750–2,200 1,300 4,100 3,300
NASEM, 19731 1,750–2,200 1,300 4,100 3,300
NASEM, 19791 1,750–2,200 1,300 4,000 2,000
NASEM, 1988 1,750–2,200 1,300 4,000 2,000
NASEM, 1998 1,750–2,200 1,300 4,000 2,000
NASEM, 2012 1,750–2,200 1,300 4,000 2,000
GfE, 2006 4,000 2,200 4,000 2,300
1
Adapted from Kim and Lindemann, 2007.

to develop a feeding program that considers the specific Results from recently conducted studies show that vi-
needs of the pigs and to monitor their vitamin A status tamin supply beyond requirement estimates of NASEM
regularly. Scientific recommendations should be used as a (2012) can improve performance and carcass characteris-
general guideline rather than a strict rule. Table 5 can pro- tics in swine (Cho et al., 2017; Santos et al., 2020; Yang
vide valuable guidance for swine nutrition, but it should et al., 2020; Hinson et al., 2022; Table 7). According to
be interpreted with caution and adjusted based on indi- Leeson and Summers (2001), when practical requirements
vidual circumstances. for vitamin A are being set, the following points must be
Although additional vitamin A requirements caused by considered:
stress, infection, or illness as well as by marginal hygiene
or temperature related-stress do not belong to nutrition- 1. Genetic differences that may affect the require-
physiological requirements, these needs must still be ac- ments.
counted for under practical conditions (DLG, 2010). In 2. Differences in bioavailability between vitamin A
Table 6, it is evident that average industrial vitamin A supplements.
fortification levels are significantly higher than those set 3. Vitamin A loss through oxidation (e.g., due to cat-
by NASEM. alytic effects of trace minerals, peroxidized polyun-
A growing body of evidence is strongly suggestive of an saturated fats, and thermal treatment of the feed).
increased need for vitamin A in swine compared with the 4. Destruction of vitamin A in the gastrointestinal
norms of GfE (2006) or NASEM (2012). The majority of tract by a variety of factors.
data used to determine current vitamin A recommenda- 5. Lower absorption in the small intestine due to in-
tions for pigs was generated in the 1950s and 1960s (Kim ternal parasites, mycotoxins, or an insufficient fat
and Lindemann, 2007). Modern pig breeds, which have content in the feed that impairs the absorption of
been genetically improved to achieve fast growth and lean fat-soluble vitamins, for example.
meat deposition, show increased metabolic turnover and 6. Insufficient protein or lipid status in the animal
oxidative stress equivalent to a marathon runner (Lau- that impairs formation of β-lipoproteins and reti-
ridsen, 2016). Because of inflated redox imbalance, the nol-binding protein for vitamin A transport.
requirement for fat-soluble antioxidant vitamins such as 7. Greater vitamin A requirements due to illness or
retinol and α-tocopherol should be higher. Furthermore, any other kind of stress.
the conversion of high-quality feed into weight gain has
significantly improved over the last 40 to 50 years in swine. When it comes to maximizing animals’ productivity, it
As a result of lower feed per unit of gain today, intake of vi- is vital to optimize feed intake and provide optimal di-
tamins per unit of gain is also lower than 40 to 50 years ago etary requirements at all stages of production. Vitamin
at the same level of supplementation. However, it is highly A requirements set by pig breeding companies can serve
unlikely that vitamin A utilization or metabolic pathway as guides for practical feeding. These recommendations
progressed with genetic selection in swine. Thus, consider- are built upon scientific publications and data, probably
able research efforts need to be expended into the current also taking into account the incorporation of a sufficient
retinol requirements in all production stages in pigs. safety margin for storage and processing losses. Table 8
Table 6. Average industrial supplementation levels of vitamin A (in IU/kg of feed) in the United States, China, Brazil, and Denmark (sources: Flohr et al., 2016 [USA 1];
Dalto and da Silva, 2020; Yang et al., 2021; Faccin et al., 2023 [USA 2]; Kjeldsen, 2023)1

USA 1 USA 2 Brazil Denmark China

Ratio to Ratio to Ratio to Ratio to Ratio to

Diet Average NASEM Average NASEM Average NASEM Average NASEM Average NASEM
Creep diets (weaning to 8 kg) 10,280 4.7
Creep diets (3 to 20 d of age) 11,368 5.2
Weaning diet (6.5 kg until wk 12 of 12,000 5.4
age)
Nursery diet 1 (weaning to 7 kg) 10,600 4.8 9,418 4.3
Nursery diets I (21 to 35 d of age) 11,037 5.0
Early nursery diets (8 to 15 kg) 10,186 4.6
Nursery diets II (36 to 49 d of age) 10,966 5.0
Nursery diet 2 (7 to 11 kg) 10,274 4.7 8,625 3.9
Weaners (9 to 20 kg) 5,649, 11,5002 3.3, 6.72
Late nursery diets (15 to 25 kg) 9,690 5.5
Nursery diets III (50 to 70 d of age) 10,014 5.8
Nursery diet 3 (11 to 23 kg) 8,868 5.1 5,951 3.4
Growing diets (71 to 120 d of age) 6,186 4.8
Growing diets (25 to 60 kg) 4,946 3.8
Finishing diets (121 d of age to 5,092 3.9
slaughter)
Early finishing diets (23 to 55 kg) 5,643 4.3 3,981 3.1
Early finishing diets (60 to 90 kg) 4,656 3.6
Mid-finishing diets 4,842 3.7 3,394 2.61
Finishers (30–100 kg) 5,000, 4,2003 3.8, 3.23
Late finishing diets (90 kg to market) 4,826 3.7
Late finishing diets (100 kg to market) 4,187 3.2 2,874 2.2
Gilt-development diets 9,405 2.4 8,497 2.1 10,273 2.6 6,640 5.1
Breeding stock 8,400 2.1
Shastak and Pelletier: Vitamin A supply in swine production

Gestation diets 10,362 2.6 9,646 2.4 8,400 2.1 7,721 1.9
Transition diets 7,800 2.0,4 3.95
Lactation diets 10,404 5.2 9,646 4.8 10,930 5.5 5,457 2.7
Boar diets 11,249 2.8 10,410 2.6 11,256 2.8 7,071 1.8
1
“Ratio to NASEM” values represent average supplementation rates as a proportion to total dietary vitamin requirements from NASEM, 2012.
2
Low-protein diet.
3
Diet with 200% phytase (double dose of phytase; e.g., 500 FTU of Natuphos = 100%, 1,000 FTU of Natuphos = 200% based on Danish nutritional standards);
4
Late gestation.
5
Early lactation.
299
300 Nutrition

provides the recommendations for vitamin A from various

pig breeding companies.

Commercial
Ultimately, it is crucial to consider the local regulatory
Hinson et al. (2022)

0.702
Table 7. Effects of increasing dietary supplementation of fat- and water-soluble vitamins (NASEM or commercial levels) on growth performance of growing swine

requirements and conditions of use when applying retinyl

23.02
4102
5902
esters, particularly in swine production. To ensure safe
and appropriate use of vitamin A in pig farming, it is im-
portant to adhere to specific guidelines that limit the sup-
NASEM

plemental levels of vitamin A to the recommended intake

0.69
21.0
levels as established by relevant authorities such as Euro-

360
530

Linear response for basal diets supplemented with 0.00%, 0.05% (NASEM level), 0.10%, 0.25%, and 0.50% (commercial level) vitamin premix.
pean Commission in the European Union, the Canadian
Food Inspection Agency in Canada, the Australian Pesti-
cides and Veterinary Medicines Authority in Australia, or
Commercial

the Ministry of Agriculture and Rural Affairs in China.

0.642
Yang et al. (2020)

Therefore, any use of vitamin A in swine nutrition must

3642
568

always be carried out within the framework of legal provi-

sions and guidelines to prevent over-supplementation and
potential negative effects on animal health and welfare.
NASEM

Adhering to these guidelines is essential, to ensure that vi-

0.56

tamin A supplementation in pig farming is safe and effec-

320
577

tive for promoting optimal health and productivity. Due

to the strict regulation of vitamin A levels in animal feed,
the occurrence of hypervitaminosis A in swine is highly
6.3 times greater

unlikely and unheard of in practical farming or husbandry

than NASEM

scenarios, even though it can be induced in certain experi-

5.42

19.73
12.3

mental conditions (EFSA, 2008).

Santos et al. (2020)

CONCLUSIONS
Retinol is a key vitamin in pig nutrition. The plant feed-
4 times greater

stuffs used to compound swine feed do not contain relevant

than NASEM

amounts of vitamin A. Therefore, synthetic retinyl acetate

5.2
12.3
19.6

supplementation is essential to meet dietary requirement.

A positive effect of optimal supplementation and injection
of retinol on swine reproduction, performance, immunity,
and health has been demonstrated. If we know at what
stages of development greater retinol supplementation
Commercial

may be appropriate and consider the effects on other vita-

0.53
Cho et al. (2017)

mins, especially vitamins E and D, we can design balanced

1
19.1
4141
7831

pig diets that will maximize performance under specific

conditions. Further investigation of retinol’s interaction
with other fat-soluble bioactive substances is needed. Ad-
ditionally, we recommend that the recommendations for
NASEM

0.58
17.4

vitamin A made by scientific committees be updated to

352
613

reflect the current state of science regarding the functions

of vitamin A, genetics, and current methods used in pig
farming.
P < 0.05 for ADWG in this phase.
BW in nursery phase 1 (d 21), kg
BW in nursery phase 2 (d 49), kg
BW in nursery phase 1 (d 63), kg

ACKNOWLEDGMENTS
BW at d 28 postweaning, kg

BW at d 40 postweaning, kg

Ute Obermueller-Jevic (BASF SE, Nutrition & Health

Division, 67063 Ludwigshafen am Rhein, Germany) is
thanked for constructive comments on the draft of the
manuscript. This work received no external funding.
Because no human or animal subjects were used, this
P < 0.05.
ADFI, g/d
ADG, g/d

ADG, g/d
ADFI, kg

analysis did not require approval by an Institutional

Study

Animal Care and Use Committee or Institutional Re-

G:F

G:F

view Board.
1

3
 Shastak and Pelletier: Vitamin A supply in swine production 301

Table 8. Vitamin A recommendations for some pig breeds1

Vitamin A recommendation, IU/kg of feed

Gestating Lactating
Breed or Finishing gilts gilts gilts
organization Piglets and barrows and sows and sows Source
DanBred 8,8001 4,0322 10,000 10,000 DanBred, 2020, Nutrient Specifications
PIC 11,000 5,510–6,600 9,920 9,920 PIC, 2016, Nutrient Specification Manual
Genesus 12,000 4,188–5,510 12,000 12,000 Genesus, 2017, Genesus Nutrient
Specifications
Topigs Min. 16,000 7,000–10,000 10,000 12,000 Topigs Norsvin, 2016, Topigs Norsvin
Manual Topigs 20
Bundes Hybrid 16,000 5,200–6,500 12,000 12,000 BHZP, 2016
Zucht Programm
Hypor 8,000 8,000 12,000 14,000 Hypor, 2017a,b
1
The amounts of supplemental vitamin A in feed shall not exceed the local regulatory maximum.
2
Recalculated based on the recommendations for “amount of retinol per MJ ME.”

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