This paper reviews the effects of extended lactation (EXT) as a strategy in dairy cattle on milk production and persistency, reproduction, milk quality, lifetime performance of the cow and finally the economic effects on herd and farm levels as well as the impact on emission of greenhouse gas at product level. Primiparous cows are able to produce equal or more milk per feeding day during EXT compared with a standard 305-d lactation, whereas results for multiparous cows are inconsistent. Cows managed for EXT can achieve a higher lifetime production while delivering milk with unchanged or improved quality properties. Delaying insemination enhances mounting behaviour and allows insemination after the cow’s energy balance has become positive. However, in most cases EXT has no effect or a non-significant positive effect on reproduction. The EXT strategy sets off a cascade of effects at herd and farm level. Thus, the EXT strategy leads to fewer calvings and thereby expected fewer diseases, fewer replacement heifers and fewer dry days per cow per year. The optimal lifetime scenario for milk production was modelled to be an EXT of 16 months for first parity cows followed by an EXT of 10 months for later lactations. Modelling studies of herd dynamics indicate a positive effect of EXT on lifetime efficiency (milk per dry matter intake), mainly originating from benefits of EXT on daily milk yield in primiparous cows and the reduced number of replacement heifers. Consequently, EXT also leads to reduced total meat production at herd level. For the farmer, EXT can give the same economic return as a traditional lactation period. At farm level, EXT can contribute to a reduction in the environmental impact of dairy production, mainly as a consequence of the reduced production of beef. A wider dissemination of the EXT concept will be supported by methods to predict which cows may be most suitable for EXT, and clarification of how milking frequency and feeding strategy through the lactation can be organised to support milk yield and an appropriate body condition at the next calving.

Improvements in feed intake of dairy cows entering the early lactation period potentially decrease the risk of metabolic disorders, but before developing approaches targeting the intake level, mechanisms controlling and dysregulating energy balance and feed intake need to be understood. This review focuses on different inflammatory pathways interfering with the neuroendocrine system regulating feed intake of periparturient dairy cows. Subacute inflammation in various peripheral organs often occurs shortly before or after calving and is associated with increased pro-inflammatory cytokine levels. These cytokines are released into the circulation and sensed by neurons located in the hypothalamus, the key brain region regulating energy balance, to signal reduction in feed intake. Besides these peripheral humoral signals, glia cells in the brain may produce pro-inflammatory cytokines independent of peripheral inflammation. Preliminary results show intensive microglia activation in early lactation, suggesting their involvement in hypothalamic inflammation and the control of feed intake of dairy cows. On the other hand, pro-inflammatory cytokine-induced activation of the vagus nerve transmits signalling to the brain, but this pathway seems not exclusively necessary to signal feed intake reduction. Yet, less studied in dairy cows so far, the endocannabinoid system links inflammation and the hypothalamic control of feed intake. Distinct endocannabinoids exert anti-inflammatory action but also stimulate the posttranslational cleavage of neuronal proopiomelanocortin towards β-endorphin, an orexigen promoting feed intake. Plasma endocannabinoid concentrations and hypothalamic β-endorphin levels increase from late pregnancy to early lactation, but less is known about the regulation of the hypothalamic endocannabinoid system during the periparturient period of dairy cows. Dietary fatty acids may modulate the formation of endocannabinoids, which opens new avenues to improve metabolic health and immune status of dairy cows.

The increasing lactational performance of dairy cows over the last few decades is closely related to higher nutritional requirements. The decrease in dry matter intake during the peripartal period results in a considerable mobilisation of body tissues (mainly fat reserves and muscle mass) to compensate for the prevailing lack of energy and nutrients. Despite the activation of adaptive mechanisms to mobilise nutrients from body tissues for maintenance and milk production, the increased metabolic load is still a risk factor for animal health. The prevalence of production diseases, particularly subclinical ketosis is high in the early lactation period. Increased β-hydroxybutyrate (BHB) concentrations further depress gluconeogenesis, feed intake and the immune system. Despite a variety of adaptation responses to nutrient and energy deficit that exists among dairy cows, an early and non-invasive detection of developing metabolic disorders in milk samples would be useful. The frequent and regular milking process of dairy cows creates the ability to obtain samples at any stage of lactation. Routine identification of biomarkers accurately characterising the physiological status of an animal is crucial for decisive strategies. The present overview recapitulates established markers measured in milk that are associated with metabolic health of dairy cows. Specifically, measurements of milk fat, protein, lactose and urea concentrations are evaluated. Changes in the ratio of milk fat to protein may indicate an increased risk for rumen acidosis and ketosis. The costly determination of individual fatty acids in milk creates barriers for grouping of fatty acids into saturated, mono- and polyunsaturated fatty acids. Novel approaches include the potential of mid-IR (MIR) based predictions of BHB and acetone in milk, although the latter are not directly measured, but only estimated via indirect associations of concomitantly altered milk composition during (sub)clinical ketosis. Although MIR-based ketone body concentrations in milk are not suitable to monitor the metabolic status of the individual cow, they provide an estimate of the overall herd or specific groups of animals earlier in a particular stage of lactation. Management decisions can be made earlier and animal health status improved by adjusting diet composition.

In cattle artificial insemination plays not only a vital role in the successful establishment of pregnancy, which is a prerequisite for initiation of the subsequent lactation, but also in accelerating genetic improvement and facilitating the distribution of semen from genetically elite sires. The latter has been greatly facilitated by the ability to successfully cryopreserve semen. The objective of an insemination is to ensure that there is an adequate reservoir of competent, capacitated, motile sperm in the caudal region of the oviductal isthmus, the site of the main sperm reservoir in the cow, at the time of ovulation to ensure fertilisation. Handling of semen, particularly the 0.25 ml straw, is critically important. Thawed semen needs to be protected from cold and heat shocks and inseminated within 6 to 8 min of thawing. Uterine horn insemination give a modest improvement in conceptions rates particularly in situations where conception rates are low following uterine body inseminations. Most of the studies that evaluated heterospermic insemination were conducted on fresh semen only, and many lacked adequate replication. Consequently, it is difficult to deduce if there are real benefits from using heterospermic semen. While the interval from oestrous onset to time of ovulation would appear to be similar for cows and heifers at about 28 h there is huge variation (standard deviations of 5 to 6 h) around this average. While best conception rates are achieved when cows are inseminated from mid oestrus to a few hours after the onset of oestrus, this is difficult to achieve in practice. There is emerging evidence that having one insemination time, when all cows requiring insemination in the herd on that day are inseminated, does not compromise fertility provided insemination technique is good and the semen used is of high fertility.

The rumen contains a great diversity of prokaryotic and eukaryotic microorganisms that allow the ruminant to utilize ligno-cellulose material and to convert non-protein nitrogen into microbial protein to obtain energy and amino acids. However, rumen fermentation also has potential deleterious consequences associated with the emissions of greenhouse gases, excessive nitrogen excreted in manure and may also adversely influence the nutritional value of ruminant products. While several strategies for optimizing the energy and nitrogen use by ruminants have been suggested, a better understanding of the key microorganisms involved and their activities is essential to manipulate rumen processes successfully. Diet is the most obvious factor influencing the rumen microbiome and fermentation. Among dietary interventions, the ban of antimicrobial growth promoters in animal production systems has led to an increasing interest in the use of plant extracts to manipulate the rumen. Plant extracts (e.g. saponins, polyphenol compounds, essential oils) have shown potential to decrease methane emissions and improve the efficiency of nitrogen utilization; however, there are limitations such as inconsistency, transient and adverse effects for their use as feed additives for ruminants. It has been proved that the host animal may also influence the rumen microbial population both as a heritable trait and through the effect of early-life nutrition on microbial population structure and function in adult ruminants. Recent developments have allowed phylogenetic information to be upscaled to metabolic information; however, research effort on cultivation of microorganisms for an in-depth study and characterization is needed. The introduction and integration of metagenomic, transcriptomic, proteomic and metabolomic techniques is offering the greatest potential of reaching a truly systems-level understanding of the rumen; studies have been focused on the prokaryotic population and a broader approach needs to be considered.

The use of a proteomic approach to investigate changes in the milk proteome is growing and has parralleled the increasing technological developments in proteomics moving from early investigation using a gel-based two-dimensional separation approach to more quantitative method of current focus applying chromatography and mass spectrometry. Proteomic approaches to investigate lactational performance have made substantial findings especially in the alterations in lactation during mastitis. An experimental model of Streptococcus uberis infection of the mammary gland has been used as a means to determine change not only in the milk proteome, but also in the peptidome and in the metabolome caused by the infection. Examination of the peptidome, that is the peptides of less than 25 kDa in molecular weight, demonstrated an increase in small peptides most of which were casein degradation products but also included small bioactive peptides such as mammary-associated serum amyloid A3 (MSAA3). The peptidome has also been shown to differ depending on the causative bacteria of naturally occuring mastitis. The use of a non-gel-based relative quantitative proteomic methodology has revealed major changes in the protein component of milk in mastitis. The S. uberis infection lead to increases in the concentrations of proteins such as cathelicidins, haptoglobin, MSAA3 and decreases milk content of proteins such as xanthine oxidase, butyrophilin and β-1,4-galactosyltransferase. Analysis of all protein change data identified the acute phase, coagulation and complement pathways as well as proteins related to bile acid metabolism as being most modified. Examination of the small molecular weight organic molecules of milk using a metabolomic approach identified an increase in the content in milk during mastitis of bile acids such as taurochenodeoxycholic acid. Notable changes were also found in metabolites responding to infection of the mammary gland. Carbohydrate and nucleic acid metabolites were reduced, whereas lipid and nitrogen containing metabolites were increased. The latter included increases in amino acids along with di and tri peptides, likely to be the result of casein degradation. The use of proteomics and other omic technology is in its infancy in investigation of lactational parameters, but can already provide additional insight into the changes involved in disease and will have further value in physiological and nutritional investigation of lactation.

It is approaching three decades since the first public evidence of sex-sorting of semen. The technology has progressed considerably since then with a number of institutions and researchers collaborating to eventually bring this to application. The technical challenges have been quite substantial and in the early years the application was limited to only heifer inseminations. Comparable fertility of sex-sorted semen with conventional semen has been an aspirational benchmark for the industry for many years. Significant investment in research in the primary biology of sex-sorted sperm and associated sorting equipment ensured steady progress over the years and current methods particularly the new SexedULTRA-4M™ seems to have now mostly bridged this fertility gap. The dairy and beef industry have adopted this technology quite rapidly. Other animal industries are progressively testing it for application in their specific niches and environments. The current state of the art in the fundamentals of sex-sorting, the biology of the process as well as new developments in machinery are described in this review.

Small ruminants not only differ on mammary gland anatomy, milk’s properties and the amount of milk yielded comparable to those of dairy cattle, but also on the milking routine strategies and machine milking settings to maximize daily milk secretion. The udder compartment is proportionally larger in dairy sheep and goats, which requires modifications in the milking machine settings, milking procedures and allows the use of different milking strategies as they better tolerate extension of milking intervals. Depending on the breed, cisternal milk in goats varies from 70% to 90%, whereas in dairy sheep it varies from 50% to 78% of the total gland capacity. This explains why these species are commonly milked without pre-milking teat preparation, while in goats it is applied only in cases of high prevalence of intramammary infection in the herd. Recent French researchers observed that 40% of the goats presented an unbalanced udder as well as unbalanced morphology (21% to 30%) and functional milk flow (around 10% to 20% more) which could induce overmilking. In dairy sheep, selection for higher milk production increases teat angle insertion. Thus, to increase machine milk fraction, it is recommended to use either the ‘Sagi hook’ as an alternative for lifting up the ‘pendulous’ udder during milking or to perform machine stripping. There are three cluster removal strategies for small ruminants: manual, timed and milk flow driven automatic cluster removal (ACR). Automatic cluster removal reduces overmilking, improves teat condition, enables labour saving and provides a consistent milking routine in small ruminants. There are three to five main milk flow profiles in ewes and goats, which result in curves with one or two peaks (or plateau) and different patterns of the milk flow decreasing phase due to the degree of mammary gland imbalance and teat characteristics. When taking into account our current knowledge, ACR recommended take-off settings for goats are: 200 g/min+10 s delay time (DT) for a long decreasing phase or two plateau curves and 500 g/min+5 s DT for a short decreasing phase and one plateau curve. The ACR take-off settings for ewes are: 150 g/min +10 s DT for long decreasing phase and 200 g /min+5 s DT for a short decreasing phase. This review is intended to be useful for scientists and producers seeking basic knowledge of milking routines and cluster detachment settings for parlour performance and milk quality.

Making dairy farming more cost-effective and reducing nitrogen environmental pollution could be reached through a reduced input of dietary protein, provided productivity is not compromised. This could be achieved through balancing dairy rations for essential amino acids (EAA) rather than their aggregate, the metabolizable protein (MP). This review revisits the estimations of the major true protein secretions in dairy cows, milk protein yield (MPY), metabolic fecal protein (MFP), endogenous urinary loss and scurf and associated AA composition. The combined efficiency with which MP (EffMP) or EAA (EffAA) is used to support protein secretions is calculated as the sum of true protein secretions (MPY + MFP + scurf) divided by the net supply (adjusted to remove the endogenous urinary excretion: MPadj and AAadj). Using the proposed protein and AA secretions, EffMP and EffAA were predicted through meta-analyses (807 treatment means) and validated using an independent database (129 treatment means). The effects of MPadj or AAadj, plus digestible energy intake (DEI), days in milk (DIM) and parity (primiparous v. multiparous), were significant in all models. Models using (MPadj, MPadj × MPadj, DEI and DEI × DEI) or (MPadj/DEI and MPadj/DEI × MPadj/DEI) had similar corrected Akaike’s information criterion, but the model using MPadj/DEI performed better in the validation database. A model that also included this ratio was, therefore, used to fitting equations to predict EffAA. These equations predicted well EffAA in the validation database except for Arg which had a strong slope bias. Predictions of MPY from predicted EffMP based on MPadj/DEI, MPadj/DEI × MPadj/DEI, DIM and parity yielded a better fit than direct predictions of MPY based on MPadj, MPadj × MPadj, DEI, DIM and parity. Predictions of MPY based on each EffAA yielded fairly similar results among AA. It is proposed to ponder the mean of MPY predictions obtained from each EffAA by the lowest prediction to retain the potential limitation from AA with the shortest supply. Overall, the revisited estimations of endogenous urinary excretion and MFP, revised AA composition of protein secretions and inclusion of a variable combined EffAA (based on AAadj/DEI, AAadj/DEI × Aadj/DEI, DIM and parity) offer the potential to improve predictions of MPY, identify which AA are potentially in short supply and, therefore, improve the AA balance of dairy rations.

Because of technical limitations, an impact of machine milking on the teat tissue cannot be avoided. The continuance of this impact during and after milking depends on a variety of factors related to the physiological regulation of milk ejection, as well as the different production systems and milking machine settings. Milking machine settings aim to achieve a high milking performance, that is, short machine-on time at a maximum of milk harvest. However, a high milking performance level is often related to an impact on the teat tissue caused by vacuum or liner compression that can lead to pathological dimensions of congestion of the tissue or hyperkeratosis as a long-term effect. Toward the end of milking a decrease of milk flow rate causes a raise of mouthpiece and teat end vacuum levels and hence an increase of the impact on the teat tissue and the risk of tissue damage. The mechanical stress by the milking machine activates a cascade of cellular mechanisms that lead to an excessive keratin growth and thickening of the keratin layer. Consequently, a complete closure of the teat canal is disabled and the risk of bacterial invasion and intramammary infection increases. Another consequence of high vacuum impact is fluid accumulation and congestion in the tissue of teat tip and teat basis because of an obstruction in venous return. The present review paper provides an overview of the available scientific information to describe the interaction between different levels and types of system vacuum, mouthpiece chamber vacuum, teat end (claw) vacuum, liner pressure, and the risk of short-term and long-term impacts on the teat tissue.

In addition to a multifactorial etiology of nutritional, social and environmental stressors, post-weaning diarrhea (PWD) in pigs is often related to infection with specific pathogens such as enterotoxigenic Escherichia coli (ETEC). In swine farming operations, the incidence of PWD is a global concern and is associated with an unbalanced gut status, resulting in poor performance and high antimicrobial consumption via prophylaxis and metaphylaxis. Increases in antimicrobial resistance are reinforcing an already-urgent need for sustainable, alternative solutions for maintaining optimal gut health in livestock. Tannin-rich plants and extracts contain bioactive compounds that could be of great interest in this respect. This review describes how the use of tannins around weaning could be beneficial for pigs, with special emphasis on the reduction of ETEC-related PWD. An overview of the broad chemical diversity of tannins is presented together with their physicochemical and biological properties, as well as how they may be metabolized in the digestive tract. The pharmacological effects exerted by tannins are summarized; more precisely, the possible mechanisms by which tannins can disrupt the different steps of the pathogenesis of ETEC-related PWD are highlighted. The factors affecting the bioactivity of tannins are also discussed, shedding light on the importance of chemical structure among different tannins.

The use of sexed semen in dairy and beef cattle production provides a number of benefits at both farm and industry levels. There is an increasing demand for dairy and beef products across the globe, which will necessitate a greater focus on improving production efficiency. In dairy farming, there is surplus production of unwanted male calves. Male dairy calves increase the risk of dystocia compared with heifer calves, and as an unwanted by-product of breeding with conventional semen, they have a low economic value. Incorporating sexed semen into the breeding programme can minimise the number of unwanted male dairy calves and reduce dystocia. Sexed semen can be used to generate herd replacements and additional heifers for herd expansion at a faster rate from within the herd, thereby minimising biosecurity risks associated with bringing in animals from different herds. Furthermore, the use of sexed semen can increase herd genetic gain compared with use of non-sorted semen. In dairy herds, a sustainable breeding strategy could combine usage of sexed semen to generate replacements only, and usage of beef semen on all dams that are not suitable for generating replacements. This results in increased genetic gain in dairy herd, increased value of beef output from the dairy herd, and reduced greenhouse gas emissions from beef. It is important to note, however, that even a small decrease in fertility of sexed semen relative to conventional semen can negate much of the economic benefit. A high fertility sexed semen product has the potential to accelerate herd expansion, minimise waste production, improve animal welfare and increase profitability compared with non-sorted conventional semen.

Assisted reproduction techniques (ARTs) provide access to early stage embryos whose analysis and assessment deliver valuable information. The handling of embryos, including the in vitro production of bovine embryos, is a rapidly evolving area which nonetheless exposes the embryos to unnatural conditions for a period of time. The Fallopian tube provides innumerable quantitative and qualitative factors, all of which guarantee the successful development of the embryo. It is well known that the Fallopian tube can be bypassed, using embryo transfer, resulting in successful implantation in the target recipient animal and the birth of calves. However, the question arises as to whether such circumvention has a negative impact on the embryo during this sensitive development period. First crosstalk between the embryo and its environment confirms mutual recognition activities and indicate bilateral effects. Nowadays, in vitro production of bovine embryos is a well-established technology. However, it is still evident that in vitro generated embryos are not qualitatively comparable to embryos obtained ex vivo. To counteract these differences, comparative studies between in vitro and ex vivo embryos are advantageous, as embryos grown in their physiological environment can provide a blueprint or gold standard against which to compare embryos produced in vitro. Attempts to harness the bovine oviduct were sometimes very invasive and did not result in wide acceptance and routine use. Long-term development and refinement of transvaginal endoscopy for accessing the bovine oviduct has meanwhile been routinely applied for research as well as in practice. Comparative studies combining in vitro development with development in the cattle oviduct revealed that the environmental conditions to which the embryo is exposed before activation of the embryonic genome can have detrimental and lasting effects on its further development. These effects are manifested as deviations in gene expression profiles and methylation signatures as well as frequency of whole chromosomal or segmental aberrations. Furthermore, it was shown that hormonal superstimulation (multiple ovulation and embryo transfer), varying progesterone concentrations as well as metabolic disorders caused by high milk production, markedly affected embryo development in the postpartum period. Assisted reproductive techniques that allow the production and handling of extra numbers of generated embryos promise to have a very high impact on scientific and practical application. Any influence on the early embryonic life, both in animals and in vitro, is accompanied by a sensitive change in embryonic activity and should be assessed in vivo on the basis of physiological conditions before being used for ART.

Artificial insemination has been a landmark procedure in improving animal agriculture over the past 150 years. The utility of artificial insemination has facilitated a rapid improvement in animal genetics across agricultural species, leading to improvements of growth, health and productivity in poultry, swine, equine and cattle species. The utility of artificial insemination, as with all assisted reproductive technologies side-steps thousands of years of evolution that has led to the development of physiological systems to ensure the transmission of genetics from generation to generation. The perceived manipulation of these physiological systems as a consequence of assisted reproduction are points of interest in which research could potentially improve the success of these technologies. Indeed, seminal fluid is either removed or substantially diluted when semen is prepared for artificial insemination in domestic species. Although seminal fluid is not a requirement for pregnancy, could the removal of seminal fluid from the ejaculate have negative consequences on reproductive outcomes that could be improved to further the economic benefit of artificial insemination? One such potential influence of seminal fluid on reproduction stems from the question; how does the allogeneic foetus survive gestation in the face of the maternal immune system? Observation of the maternal immune system during pregnancy has noted maternal immune tolerance to paternal-specific antigens; a mechanism by which the maternal immune system tolerates specific paternal antigens expressed on the foetus. In species like human or rodent, implantation occurs days after fertilisation and as such the mechanisms to establish antigen-specific tolerance must be initiated very early during pregnancy. We and others propose that these mechanisms are initiated at the time of insemination when paternal antigens are first introduced to the maternal immune system. It is unclear whether such mechanisms would also be involved in domestic species, such as cattle, where implantation occurs weeks later in gestation. A new paradigm detailing the importance of paternal–maternal communication at the time of insemination is becoming evident as it relates to maternal tolerance to foetal antigen and ultimately pregnancy success.

Millions or billions of sperm are deposited by artificial insemination or natural mating into the cow reproductive tract but only a few arrive at the site of fertilization and only one fertilizes an oocyte. The remarkable journey that successful sperm take to reach an oocyte is long and tortuous, and includes movement through viscous fluid, avoiding dead ends and hostile immune cells. The privileged collection of sperm that complete this journey must pass selection steps in the vagina, cervix, uterus, utero-tubal junction and oviduct. In many locations in the female reproductive tract, sperm interact with the epithelium and the luminal fluid, which can affect sperm motility and function. Sperm must also be tolerated by the immune system of the female for an adequate time to allow fertilization to occur. This review emphasizes literature about cattle but also includes work in other species that emphasizes critical broad concepts. Although all parts of the female reproductive tract are reviewed, particular attention is given to the sperm destination, the oviduct.

Methane (CH4) production is a ubiquitous, apparently unavoidable side effect of fermentative fibre digestion by symbiotic microbiota in mammalian herbivores. Here, a data compilation is presented of in vivo CH4 measurements in individuals of 37 mammalian herbivore species fed forage-only diets, from the literature and from hitherto unpublished measurements. In contrast to previous claims, absolute CH4 emissions scaled linearly to DM intake, and CH4 yields (per DM or gross energy intake) did not vary significantly with body mass. CH4 physiology hence cannot be construed to represent an intrinsic ruminant or herbivore body size limitation. The dataset does not support traditional dichotomies of CH4 emission intensity between ruminants and nonruminants, or between foregut and hindgut fermenters. Several rodent hindgut fermenters and nonruminant foregut fermenters emit CH4 of a magnitude as high as ruminants of similar size, intake level, digesta retention or gut capacity. By contrast, equids, macropods (kangaroos) and rabbits produce few CH4 and have low CH4 : CO2 ratios for their size, intake level, digesta retention or gut capacity, ruling out these factors as explanation for interspecific variation. These findings lead to the conclusion that still unidentified host-specific factors other than digesta retention characteristics, or the presence of rumination or a foregut, influence CH4 production. Measurements of CH4 yield per digested fibre indicate that the amount of CH4 produced during fibre digestion varies not only across but also within species, possibly pointing towards variation in microbiota functionality. Recent findings on the genetic control of microbiome composition, including methanogens, raise the question about the benefits methanogens provide for many (but apparently not to the same extent for all) species, which possibly prevented the evolution of the hosting of low-methanogenic microbiota across mammals.

Fertilization is an intricate cascade of events that irreversibly alter the participating male and female gamete and ultimately lead to the union of paternal and maternal genomes in the zygote. Fertilization starts with sperm capacitation within the oviductal sperm reservoir, followed by gamete recognition, sperm–zona pellucida interactions and sperm–oolemma adhesion and fusion, followed by sperm incorporation, oocyte activation, pronuclear development and embryo cleavage. At fertilization, bull spermatozoon loses its acrosome and plasma membrane components and contributes chromosomes, centriole, perinuclear theca proteins and regulatory RNAs to the zygote. While also incorporated in oocyte cytoplasm, structures of the sperm tail, including mitochondrial sheath, axoneme, fibrous sheath and outer dense fibers are degraded and recycled. The ability of some of these sperm contributed components to give rise to functional zygotic structures and properly induce embryonic development may vary between bulls, bearing on their reproductive performance, and on the fitness, health, fertility and production traits of their offspring. Proper functioning, recycling and remodeling of gamete structures at fertilization is aided by the ubiquitin–proteasome system (UPS), the universal substrate-specific protein recycling pathway present in bovine and other mammalian oocytes and spermatozoa. This review is focused on the aspects of UPS relevant to bovine fertilization and bull fertility.

Ruminants living in seasonal environments face a two-fold challenge during winter. The energetic cost of maintaining a high body temperature is higher at lower ambient temperatures, and this is compounded by poor availability and quality of feed. Wild ruminants acclimatize to this energetic challenge by hypothermia, that is, reduced endogenous heat production and abandoning the maintenance of a high body temperature, particularly in peripheral body parts. Further but lesser contributions to lower energy expenditure during winter are reduced foraging activity; lower heat increment of feeding; and reduced maintenance cost of size-reduced organs. Altogether, metabolic rate, estimated by the continuous measurement of heart rate, during winter is downregulated to more than half of the summer level, as is voluntary food intake, even in animals fed ad libitum. The transformation from the summer into the thrifty winter phenotype is also evident in the physiology of digestion. Microbial protein synthesis is less facilitated by diminished phosphorus secretion into the shrunk rumen during winter. In line with this result, the concentration of ammonia, the end-product of protein digestion in the rumen, peaks in rumen liquid in spring, whereas the molar proportion of acetate, an indicator of fermentation of a diet rich in fiber, peaks in winter. In contrast to reduced stimulation of growth of ruminal microbes during winter, active transport of nutrients across the intestinal epithelium is increased, resulting in more efficient exploitation of the lower amount and quality of ingested winter feed. Nevertheless, the energy balance remains negative during winter. This is compensated by using fat reserves accumulated during summer, which become a major metabolic fuel during winter.

Several structural and functional features keep bull testes 2°C to 6°C below body temperature, essential for the production of morphologically normal, motile and fertile sperm. The testicular vascular cone (TVC), located above the testis, consists of a highly coiled testicular artery surrounded by a complex network of small veins (pampiniform plexus). The TVC functions as a counter-current heat exchanger to transfer heat from the testicular artery to the testicular vein, cooling blood before it enters the testis. Bulls with increased TVC diameter or decreased distance between arterial and venous blood, have a greater percentage of morphologically normal sperm. Both the scrotum and testes are warmest at the origin of their blood supply (top of scrotum and bottom of testis), but they are cooler distal to that point. In situ, these opposing temperature gradients result in a nearly uniform testicular temperature (top to bottom), cooler than body temperature. The major source of testicular heat is blood flow, not testicular metabolism. High ambient temperatures have less deleterious effects on spermatogenesis in Bos indicus v. Bos taurus bulls; differences in TVC morphology in B. indicus bulls confer a better testicular blood supply and promote heat transfer. There is a long-standing paradigm that testes operate on the brink of hypoxia, increased testicular temperature does not increase blood flow, and the resulting hypoxia reduces morphologically normal and motile sperm following testicular hyperthermia. However, in recent studies in rams, either systemic hypoxia or increased testicular temperature increased testicular blood flow and there were sufficient increases in oxygen uptake to prevent tissue hypoxia. Therefore, effects of increased testicular temperature were attributed to testicular temperature per se and not to secondary hypoxia. There are many causes of increased testicular temperature, including high ambient temperatures, fever, increased recumbency, high-energy diets, or experimental insulation of the scrotum or the scrotal neck. It is well known that increased testicular temperatures have adverse effects on spermatogenesis. Heat affects all germ cells and all stages of spermatogenesis, with substantial increases in temperature and/or extended intervals of increased testicular temperature having the most profound effects. Increased testicular temperature has adverse effects on percentages of motile, live and morphologically normal sperm. In particular, increased testicular temperature increases the percentage of sperm with abnormal morphology, particularly head defects. Despite differences among bulls in the kind and percentage of abnormal sperm, the interval from increased testicular temperature to the emergence of specific sperm defects is consistent and predictable. Scrotal surface temperatures and structural characteristics of the testis and TVC can be assessed with IR thermography and ultrasonography, respectively.

Feeding management of the postnatal and preweaning calf has an important impact on calf growth and development during this critical period and affects the health and well-being of the calves. After birth, an immediate and sufficient colostrum supply is a prerequisite for successful calf rearing. Colostrum provides high amounts of nutrient as well as non-nutrient factors that promote the immune system and intestinal maturation of the calf. The maturation and function of the neonatal intestine enable the calf to digest and absorb the nutrients provided by colostrum and milk. Therefore, colostrum intake supports the start of anabolic processes in several tissues, stimulating postnatal body growth and organ development. After the colostrum feeding period, an intensive milk feeding protocol, that is, at least 20% of BW milk intake/day, is required to realise the calf potential for growth and organ development during the preweaning period. Insufficient milk intake delays postnatal growth and may have detrimental effects on organ development, for example, the intestine and the mammary gland. The somatotropic axis as the main postnatal endocrine regulatory system for body growth is stimulated by the intake of high amounts of colostrum and milk and indicates the promotion of anabolic metabolism in calves. The development of the forestomach is an important issue during the preweaning period in calves, and forestomach maturation is best achieved by solid feed intake. Unfortunately, intensive milk-feeding programmes compromise solid feed intake during the first weeks of life. In the more natural situation for beef calves, when milk and solid feed intake occurs at the same time, calves benefit from the high milk intake as evidenced by enhanced body growth and organ maturation without impaired forestomach development during weaning. To realise an intensive milk-feeding programme, it is recommended that the weaning process should not start too early and that solid feed intake should be at a high extent despite intensive milk feeding. A feeding concept based on intensive milk feeding prevents hunger and abnormal behaviour of the calves and fits the principles of animal welfare during preweaning calf rearing. Studies on milk performance in dairy cows indicate that feeding management during early calf rearing influences lifetime performance. Therefore, an intensive milk-feeding programme affects immediate as well as long-term performance, probably by programming metabolic pathways during the preweaning period.