1

Class Assignment (ANIM 7102)

Ali Hassan Nawaz

Student ID: 2022205048

Review
PHYSIOLOGICAL EFFECTS OF NUTRITION ON THE PUBERTY ONSET AND

SPERMATOGENESIS IN CATTLE

Abstract

Nutrition affects both production and reproduction of dairy animals. Irrational feeding and

severe metabolic disorders affect dairy cattle's productive and reproductive performance.

The essential nutrients, minerals, and vitamins needed in the optimal amount are energy

and protein and must be of utmost significance to enhancing reproduction in dairy cattle.

Minerals and vitamins cannot be ignored and must be balanced in the diet. This article

attempted to discuss the effects of nutrients on early puberty and spermatogenesis in dairy

cattle. Spermatogenesis is also a nourishment-dependent process. Spermatogenesis is a

process that produces mature male gametes, which fertilize female gametes to produce

zygotes, which are single-celled organisms. A fetus is produced as a result of cell division

and growth. The number of chromosomes must be adequately maintained throughout the

body for a healthy progeny because failure can result in various defects. Upscale

production can be achieved by ameliorating production efficiency due to high animal protein

demand worldwide. Hereditary exceptional bulls are being used on a large scale for natural

breeding or AI recognition. Using genomic markers (prioritizing male and female fertility

characters) is advantageous if bulls are selected early. However, proper management is

necessary, including the development of bull perception and the influence of nutrition. Micro
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and macronutrients in optimum quantity play their role in spermatogenesis. Malnutrition

causes adverse effects on reproductive capacity and the development of sex organs. It

decreases aldosterone emission and sperm production. Malnutrition affects the

hypothalamic-pituitary axis and significantly disturbs the production and secretion of

luteinizing and follicle-stimulating hormones.

KEYWORDS

Malnutrition; puberty; spermatogenesis; dairy cattle; hormone; macronutrients

Introduction

In animals, puberty is the stage of life at which sexual organs are fully matured, and the

animal can release mature gametes (Bo et al., 2020). In males, estradiol and androgen

pass the blood-brain barrier (BBB) and before changed to estradiol via aromatase enzyme

(Lea and England, 2019; Abeygunawardena and Dematawewa, 2004; Ciccioli et al., 2005).

Nutrition affects reproduction through changes in metabolic substrates (glucose and amino

acids) and the relative direct effects of metabolic hormones on reproductive hormones and

gonadotropins (Funston et al., 2012; Perrier et al., 2020; Nogueira, 2004). The mechanism

of nutrition on follicular growth is shown in (Fig.1). Malnourished females have a deficiency

in ovarian activity due to the low level of luteinizing hormone in a pulsatile manner (Gasser

et al., 2018; Rasby and Funston, 2016). Prepubertal growth is influenced by the amount of

nutrition and luteinizing hormone (LH) pulse in the hypothalamus Chagas et al. (2007). In

undernourished cows, a decrease in the circulating level of gonadotropins and ovarian

activity leads to infertility. In males, malnutrition affects the initiation of puberty. Reduced

FSH, LH, and testosterone levels lead to decreased spermatogenesis and fertility

(Loutchanwoot and Vortherms (2018). As puberty proceeds, there is a reduction in the

activity of negative response on luteinizing hormone-releasing hormone (LHRH) by

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estradiol which raises the level of LH hormone. The LH hormone promotes follicular growth,

which then produces estradiol. This increase in estradiol induces puberty LH surge (Hall et

al., 2009; Perry, 2016). Despite other management-related practices, nutritional regimen

and feeding system directly correlates with the onset of puberty and reduction of the age of

sexual maturity in dairy animals. It also contributes as a booster of sexual vigor and normal

regulation of spermatogenesis and thus produces spermatozoa enriched with sperm quality

and fertility parameters. Increased LH secretion during the early rising is thought to be

responsible for the beneficial effects of early feeding on postpubertal bulls. Furthermore,

early nutrition predetermines age at puberty, testis size at sexual maturity, and sperm

production potential since the negative effects of restricted food throughout early life are not

reversed by nutritional supplements during the pubertal era. IGF-I may play a role in

controlling the early rise in gonadotropins because it rises concurrently with the early rise in

gonadotropins.(Thundathil et al., 2016). In this article, we enlightened briefly about different

vital dietary nutrients and their physiological interactions with critical metabolic pathways

involved in attaining puberty and undergoing normal spermatogenesis in dairy animals

Puberty in female cattle

A cow with early puberty can give birth to a calf at an initial stage and synthesize more milk

in her lifespan. However, late maturity produces a negative outcome on the economics of a

farm. Many factors affect the age of puberty in heifers, including genetics, nutrition,

environment, and hormones. The standard stage of puberty in heifer is between 34 to 37

months, but Sahiwal breed cow heifer attains puberty at 46 months Hussain et al. (2006).

In tropical regions, the age at puberty in Bos indicus ranges from 16 to 40 months

(McDowell, 1976). Before sexual maturity, the ovary does not release adequate estradiol to

trigger the surge center of the hypothalamic axis to synthesize plenty of gonadotrophin-

releasing hormones (GnRH). A low level of estradiol hurts the hypothalamic tonic center. As
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the transformation to sexual maturity starts, the sensation of the tonic center towards

estradiol restriction decreases, allowing the hypothalamic cells to release GnRH that

triggers the ovary to synthesize a superior level of estradiol. When estradiol amount

increases, the surge center secretes a preovulatory GnRH surge, which tries to induce a

preovulatory surge of LH. As a result, ovulation occurs at the beginning of puberty. Puberty

onset in the female is shown in (Fig. 2).

Puberty in bulls

The testes, epididymis, and vas deference are part of the male reproductive system. The

primary purpose of the male reproductive system is to release gametes and hormones. The

process of production of mature spermatozoa is called spermatogenesis. It includes male

gamete development of a spermatocyte from a spermatogonium, the meiotic partition of the

spermatocyte, and the renovation of four resulting spermatids into spermatozoa. In the first

meiotic division, the cells in the prophase are primary spermatocytes. After first meiotic

division, primary spermatocytes are converted into secondary spermatocytes. The cells

rapidly progress to the second meiotic division. The product of the second meiotic division

is called spermatid. These remain sphere-shaped cells with interphase nuclei located high

in the epithelium. Meanwhile, spermatids pass through a transfiguration into spermatozoa

Cheah and Yang (2011). Spermatogenesis is the process of metamorphosis of sphere-

shaped spermatids into extended spermatozoa. During spermatogenesis, acrosome forms

flagellar apparatus. Gametes are out into the lumen of the seminiferous tubules. A single

Sertoli cell spreads from the basement to the lumen of the seminiferous tubule. Due to a

tight interchange in the middle of Sertoli cells, big particles cannot pass from the blood into

a seminiferous tubule's lumen. The purpose of the blood-testis barrier may be to avoid an

autoimmune response Cheah and Yang (2011). A study has shown that puberty begins

when >50 ×106 sperms are present in the ejaculate with >10 % motility. Infantile, pre-
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pubertal, and pubertal are the bull's three reproductive developmental stages. In the

infantile period (0–8 weeks of age), gonadotropins and testosterone are less released.

Afterward, in the pre-pubertal stage (8 to 20 weeks), there is a transitory rise in the level of

blood gonadotropin concentrations and a simultaneous increase in testosterone release

Barth et al. (2008). LH concentrations rise at 4 to 5 weeks, maximum at 12 to 16 weeks,

and formerly drop, reaching a basel level at 25 weeks Dance et al. (2015). The LH

concentrations in the pre-pubertal stage affect sexual growth and are reversely related to

puberty age. Blood FSH amount usually rises throughout the early increase and drops to

Basel level around the 25 weeks. A study described that bulls supplemented with high

nutrition diet were heavier and had a sizeable scrotal circumference (SC) and larger

testicular vascular cone diameter. It regulates testicular temperature to effectively produce

fertile spermatozoa Dance et al. (2015). The estradiol primarily causes the inhibitory effect.

As puberty begins in males, hypothalamic GnRH secretion is decreased by the inhibitory

effect of estradiol and testosterone. The increased GnRH pulse releases more LH and

FSH, which excite testis to go through the spermatogenesis process beginning the onset of

puberty Lea and England (2019), as shown in (Fig. 3).

Effect of nutrition on puberty

Concentrate feeding in bulls

Early puberty will advance the availability of semen, reduce the generation gap, and hasten

genetic improvement in young, genomically chosen, high genetic merit dairy bulls. Recent

research from (Byrne et al., 2018)showed that providing bull calves with a high plane of

nutrition in the early stages of life accelerates the age at which puberty begins by around

one month. Another study showed that (Dance et al., 2016) on Holstein-Friesian bulls have

conclusively demonstrated that, despite increasing scrotal skin thickness compared to their
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contemporaries offered a moderate plane of nutrition, an enhanced plane of nutrition up to

6 months of age has no latent impacts on measures of post-pubertal semen quality,

including pre-and post-thaw sperm motility, or on the in vitro fertilizing ability (Perrier et al.,

2020). Animals supplemented with a high concentrate diet have developed more fat

accumulation around the pampiniform plexus, which may affect the thermoregulation of the

scrotal substances decreasing epididymis sperm storage and likely sperm production

(Brown, 2007). Defects related to the morphology of spermatozoa may also occur in this

condition. In low concentrate feed (8.09 to 1.35%) or protein-free (urea-containing)

decreased the sperm-production capability or the proportion of defective gametes (Brown,

2007). In contrast, feeding 100% TDN for one year to bull calves that earlier were low low-

plane nutrition (60% TDN) from eight weeks to 46 months of age had a non-significant

effect on the development of testis, sperm concentration, and sperm production. Bulls fed

with more energy intakes had considerably decreased reproductive potential compared to

average energy diets group bulls Kumar et al. (2017).

Concentrate feeding in females

Balanced nutrition is essential for puberty onset (Heinrichs et al. (2005). At a growing age,

colostrum is essential for the calf. A good quality calf starter should contain 18% crude

protein and 3.0 Mcal metabolizable energy Kumar et al. (2017). The animals fed with green

fodder along with 20 kg concentrate reached maturity earlier (727.77 ± 44.17days) than the

control group ( 993.33 ± 68.78 days) Rafiq et al. (2008). Bhatti et al. (2007) reported that

puberty age was reduced by feeding on green fodder plus concentrate compared to the

other group fed on fodders only (Table. 1). However, adding minerals to the diet had no

such type of effect on puberty age in heifers. Another study showed that in the summer

season, feeding concentrate in addition to fodder decreased the puberty age in heifers
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Ciccioli et al. (2005). In Pakistan, animals are fed on low-quality forages and fodders only.

Still, the concentrate has CP and energy values less than the standard level for temperate

breeds. These studies provided enough knowledge to reduce the puberty age by feeding

high CP and energy level concentrate before a total mixed ration.

Vitamins and minerals feeding

Males

During the post-weaning period, both undernutrition and overnutrition can have negative

impacts. Undernutrition results in delayed puberty, and overnutrition can reduce semen

quality and body condition scoring. These conditions should be closely monitored. Excellent

quality vitamins and minerals should be included in the nutrition plan as they have many

roles in animal growth and reproductive performance Walker et al. (2009). Zinc plays a vital

role in ribonuclease activity throughout mitosis of spermatogonia and meiosis of

spermatocytes Canada and Current (1984). Low zinc intake delays testicular development

and spermatogenesis cessation. Low Zinc level in the blood affects spermatogenesis and

reduces seminiferous tubules (Table. 2) (Liu et al., 2009; Cheah and Yang, 2011).

Selenium is needed for normal spermatogenesis in the male reproductive tract. It plays an

essential part in mammalian sperm maturation, but when its concentration is increased or

decreased, it causes interference in sperm development Kehr et al. (2009). Selenium

deficient diet showed deformities in the tail, midpiece, plasma membrane, and

mitochondrial gaps. Moreover, spermatogenesis is a selenium-dependent process, and the

morphology of male reproductive structures depends on the selenium's nutritional amount.

However, organic selenium has a better result due to its higher concentration and lower

toxicity Ahsan et al. (2014). Selenium concentration is elevated during spermatogenesis.

The excellent Se absorption throughout spermatogenesis is associated with its defending

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property and is linked with enzymes, such as mitochondrial capsule protein in spermatozoa

Kaur et al. (2005). Other trace metals that act similarly in spermatogenesis are nickel (Ni)

and manganese (Mn). Previous studies have pointed out that nickel deficiency leads to

defects in reproductive function. Nickel deficiency reduces sperm production in the testis,

low sperm count in the epididymis, and sperm motility. Manganese is a stimulator of

puberty growth (Lukac et al., 2014; Prestifilippo et al., 2008). Initial histological studies have

shown that vitamin deficiency in nutrient intake affects male reproductive structures (Table.

3)( Anshuman Kumar et al., 2016; Cheah and Yang, 2011).

Females

Minerals play a vital role in early maturity, essential for proper functioning reproductive

hormones and enzymes. An inadequate supply of minerals leads to anestrus Bodla et al.

(2017). Buffalo heifers raised underneath the pasture system reach puberty before than

raised under the stall feeding system. Also, minerals administration might be an approach

to effect estrus that crosses the average age at puberty onset Vierboom et al. (2003). One

study showed high zinc requirements during parturition and lactation; the body absorbs and

deposits zinc more efficiently during the third trimester of pregnancy. Zn and Cu are

involved in sexual development, reproductive growth, and maintenance of ruminants.

Studies showed that serum binding retinol (precursor of vitamin A) continuously increased

in puberty, indicating that vitamin is necessary for puberty onset. Vitamin D deficiency may

result in delayed onset of estrus and ovarian inactivity Kumar et al. (2016)

Protein diet

Male

Low energy and low protein diet cause infertility, which leads to decreased appetite.

Moreover, research showed that a high intake of crude protein changes the concentration
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of different minerals in seminiferous tubules. However, in males, nutrition is also directly

proportional to early reproductive development. In a research bull in 2 groups, one group

was fed a high dietary plane (grain), and the other was fed on a moderate nutritional plane

(roughages). Results showed that bulls of the roughages group generally reduced scrotal

superficial area heat and minor semen quality by 1% motility, normal morphology, head

defects, tail faults, etc. Bourgon et al. (2018) research was conducted in which 26 Holstein

bull calves were divided into three groups at almost one week of age to obtain a low,

medium, and great diet based on the energy level and protein from 2 to 31 st week of age.

Bull calves nourished with a significant nutrition intake reach puberty prior and have

superior testes than those nourished with a stumpy nutrition diet. It contains 20 % CP

(crude protein) medium nutrition diet contains 17% CP. A low-nutrition diet contains 12.2%

CP. The high nutrition group was fed ad libitum. The exact amount of feed was presented to

low and medium nutrition groups. Calves were fed milk 4,6,8 L/d in the small, medium, and

high nutrition diet, respectively, from 2 to 8 weeks of age; after that, the feed was offered to

the high nutrition group Dance et al. (2015).

Female

Nutrition affects the production and release of GnRH, FSH, LH, and GH due to its action on

the hypothalamus and anterior pituitary. Ovarian follicular development and steroid

production are influenced by nutrition. Acute /chronic dietary limitation results in a gradual

decline in the growth rate of the dominant follicle (Pradesh, 2016). The study was

conducted by Pedroso et al. (2018) to evaluate the effect of rumen-undegradable protein

(RUP) on feed consumption, performance, mammary gland growth, and carcass quality of

Holstein heifers in prepubertal and pubertal dairy heifers. Four diets with different RUP

concentrations (38, 44, 51, and 57% crude protein) were used. Results showed that RUP
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amounts had no effects on mammary gland composition. In contrast, pre-pubertal heifers

had greater pixel standards than pubertal heifers, indicating higher contents of fat rather

than protein in the mammary glands of pre-pubertal heifers. In a study by Manthey et al.

(2017), they checked the effect of increasing the concentration of distillers dried grains

(DDGS) by replacing the forage on the dietary fatty acid, metabolic hormones profile, and

onset of puberty in heifers. The increasing concentration of distillers dry grains (DDGS)

were 30% DDGS, 40% DDGS, and 50% DDGS, with a diet having grass hay and 1.5%

mineral mixture. Results showed that treatments had no significant effects on Body Weight

and puberty. In another study, authors determined the growth rate, body composition, and

age of puberty in Angus and Brangus heifers by supplementing soybean meal (SBM), a

source of rumen-degradable protein (RDP), with a dried distillers grain (DDG) and Bermuda

grass round bale silage (RBS) diet. Treatments were on day one RBS + DDG at 0.75%

BW; RBS + DDG at 0.75% BW + SBM at 7.5% of DDG; or RBS + DDG at 0.75% BW +

SBM at 15% of DDG for 140 day. In the end, soybean meal at also 7.5 or 15% of the total

DDG supplement had no positive results on growth rate, body composition, or age of

puberty in heifer Alava et al. (2017).

Hormones

Male

Bulls receiving GnRH had larger testicular size and attained puberty earlier than the control

group due to increased LH concentration. Bulls treated with LH releasing hormone

positively impacted testicular function, inducing a rise in Sertoli and germ cells. A high level

of LH causes early puberty, testicular enlargement, and increased spermatogenesis cycle

Hull and Harvey (2014). Upon histological evaluation, bulls treated with FSH hormone

showed more Sertoli and sperm cells ( Harstine, 2018).

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Female

A cow's fertility depends upon the reproductive hormonal profile before and after

insemination. Higher concentrations of progesterone in the blood throughout the luteal

phase of the estrous cycle preceding insemination are related to higher conception rates.

Sonderman and Larson (2010) reported that administering a gonadotropin-releasing

hormone (GnRH) analog to repeat breeder cows on day 12 of the estrous cycle resulted in

a higher embryo recovery rate and increased the percentage of normal embryos compared

to untreated control cows. The authors speculated that such a treatment ten days before

the following ovulation might have resulted in improved follicular recruitment for ovulation

leading to increased progesterone secretion by the corpus luteum Sonderman and Larson

(2010). Exogenous progestin is considered suitable for anestrous postpartum cows. The

growth of precise intravaginal progesterone-releasing strategies or an intravaginal

progesterone insert (IPI) speeds up the restoration of ovarian follicular activity. At the end of

IPI treatment, hormones such as equine chorionic gonadotropin (Fig.4), estradiol, or PGF2α

maximize follicular activity Peter et al. (2009). Numerous arrangements of GnRH analogs

and PGF2 α have been used to trigger ovarian cyclicity.

Macronutrients that affect spermatogenesis

Lipid feeding

The plasma membrane of sperm is dissimilar from other cell membranes. It includes a

more considerable amount of polyunsaturated fatty acids (PUFA), mostly diPUFA

(phospholipids esterified with two PUFA) Wassall and Stillwell (2009). The membrane

fluidity and flexibility are due to PUFA Mandal et al. (2014). Membrane lipid composition

has been associated with their particular purposes because it encourages the formation of

micro-domains with additional flexibility, fusogenicity, and permeability appearances

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Wassall and Stillwell (2009). Lipid compositions and associated antioxidant capabilities of

spermatozoa and seminal plasma from bulls were observed at the beginning, middle, and

end of their reproductive period. The reduction in absorption and motility of spermatozoa

related to aging was accompanied by a significant decrease in lipid levels within the

seminal plasma. This change in lipid concentration was accompanied by an increase in the

amount of phospholipid.

By comparison, the number of phospholipids in the spermatozoa was significantly

decreased Melville et al. (2012). In another study, Gürler et al. (2015) examined the special

effects of feeding alpha-linolenic acid (ALA) on fatty acid arrangement and superiority of

bovine sperm and the antioxidative ability of seminal plasma. Feeding ALA increased (P <

0.05) alpha-linolenic acid (ALA)content in bulls; however, no variation was included in bulls.

PMAI was better after cryopreservation in ALA and PA bulls during the experiment (P <

0.005). The LPO of the direct sperm after thawing did not change in the ALA group during

the study but decreased in the (palmitic acid PA bulls) PA group (P < 0.006). After 3 hours

of incubation, LPO increased in the ALA group (P < 0.02), while there was no difference in

LPO between groups.

In summary, the intake of unsaturated or polyunsaturated fatty acids affects the level of

antioxidants in the seminal plasma, Gürler et al. (2015). Although docosahexaenoic acid

content in sperm membranes is only increased in ALA bulls, saturated fatty acids and

polyunsaturated fatty acids have a positive and extraordinary effect on the quality of

cryopreserved bovine sperm. Testis contains a unique content of polyunsaturated fatty

acids. Spermatozoa are rich in lipids (70% of total) Conquer et al. (2000). Fats have an

essential part in maintaining sperm cell viability, maturity, fruitfulness, and functional

deficiency of PUFAs polyunsaturated fatty acids cause testicular degeneration and infertility

Mandal et al. (2014).

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Amino acids

Arginine (Arg) plays an essential role in spermatogenesis. It is an organic forerunner in the

production of spermidine and spermine and is crucial for sperm motility. L-carnitine is

involved in energy metabolism and promotes sperm motility, maturation, and sperm

development by providing voluntary energy to sperm Blackman et al. (2004). In ruminants,

if metabolizable energy is not specified, degradation of dietary protein by the rumen

microorganisms offers the animal a sensibly continuous amino-acid source causing slow

development or early pregnancy. By post-ruminal digestion of undegraded proteins, the

animal immovably provides the animal with crucial other amino acids necessary to promote

rapid development. Although some amino acid seems crucial for spermatogenesis ( Cheah

and Yang, 2011; Brown, 2007), grazing ruminants usually do not encounter shortages of

these materials unless the protein content of the pasture is low. Young cows could produce

all the essential and non-essential amino acids needed for fertility, and even in a protein-

free diet, urea is the only source of dietary nitrogen.

On the other hand, monogastric animals, such as pigs, rely heavily on dietary protein for

amino acids. However, since these animals are concentrated, most are hand-fed with a

proprietary feed mixture and should not be protein-deficient (Brown, 2007). Additionally,

antagonists of neuroexcitatory amino acids have been found to block GnRH release from

the hypothalamus and inhibit LH release. In contrast, pretreatment with GnRH receptor

antagonists abolished the ability of N-methyl-D and L-aspartate to stimulate LH release

Gürler et al. (2015). Therefore, there is a need to continuously supply dietary amino acids,

vitamins, and energy for essential tissue maintenance, body temperature regulation, and

locomotors requirement for foraging.

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Protein

Protein deficiency or low protein level in the feed may be associated with retarded

spermatogenesis, low sperm count, and inability to produce mature spermatozoa. Huang

and Sha (2011). Maximize bull breeding by providing adequate calf nutrition and proper

nutrition after weaning. The outcomes of experiments recommended that a target for an

average daily gain throughout calf-hood should be >1.2 kg/d Barth et al. (2008). In another

study, two experiments showed the effects of low protein diets on young beef cattle's

growth and reproductive physiology. In both studies, low protein diets severely reduced

feed intake and produced significant weight loss. Bulls that lack protein produces a minimal

amount of semen. In both studies, by the end of the experimental period, for calves lacking

protein, the total amount of sperm per ejaculation was less, and the percentage of motile

sperm was different. It was not affected by the treatment Bailoni et al. (2010).

Conclusion

In cows and bulls, nutrition and reproduction are closely connected. In different phases,

dairy cattle's spermatogenesis and reproductive ability can be improved by balanced

nutrition. Due to the rising demand for animal protein around the world, upscale production

can be accomplished by improving production efficiency.

For natural breeding or AI identification, hereditary exceptional bulls are widely used. If bulls

are chosen early, using genetic markers which prioritize male and female fertility traits is

favorable. However, correct management is required, which includes taking into account

how a bull perceives itself and the impact of nutrition. The performance of animals and the

security of the world's food supply will be significantly improved by the selection of

genetically superior bulls, proper management of their reproductive development, rigorous

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laboratory assays to ensure semen quality, and effective use of such semen (conventional

or sexed).

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