Animal 18 (2024) 101141

Contents lists available at ScienceDirect

Animal
The international journal of animal biosciences

Review: Opportunities and challenges for the genetic selection of dairy

calf disease traits
C. Lynch a, E.M. Leishman a,b, F. Miglior a,c, D. Kelton d, F.S. Schenkel a, C.F. Baes a,e,⇑
a
Centre for the Genetic Improvement of Livestock, Department of Animal Biosciences, University of Guelph, 50 Stone Road East, Guelph, ON, Canada
b
Centre for Nutrition Modelling, Department of Animal Biosciences, University of Guelph, 50 Stone Road East, Guelph, ON, Canada
c
Lactanet Canada, Guelph, ON N1K-1E5, Canada
d
Department of Population Medicine, University of Guelph, Ontario N1G-2W1, Canada
e
Institute of Genetics, Department of Clinical Research and Veterinary Public Health, University of Bern, Bern 3001, Switzerland

a r t i c l e i n f o a b s t r a c t

Article history: Interest in dairy cow health continues to grow as we better understand health’s relationship with produc-
Received 8 October 2023 tion potential and animal welfare. Over the past decade, efforts have been made to incorporate health
Revised 14 March 2024 traits into national genetic evaluations. However, they have focused on the mature cow, with calf health
Accepted 15 March 2024
largely being neglected. Diarrhoea and respiratory disease comprise the main illnesses with regard to calf
Available online 22 March 2024
health. Conventional methods to control calf disease involve early separation of calves from the dam and
housing calves individually. However, public concern regarding these methods, and growing evidence
Keywords:
that these methods may negatively impact calf development, mean the dairy industry may move away
Dairy calves
Diarrhoea
from these practices. Genetic selection may be a promising tool to address these major disease issues.
Genetic selection In this review, we examined current literature for enhancing calf health through genetics and discussed
Respiratory disease alternative approaches to improve calf health via the use of epidemiological modelling approaches, and
Calf health the potential of indirectly selecting for improved calf health through improving colostrum quality.
Heritability estimates on the observed scale for diarrhoea ranged from 0.03 to 0.20, while for respiratory
disease, estimates ranged from 0.02 to 0.24. The breadth in these ranges is due, at least in part, to differ-
ences in disease prevalence, population structure, data editing and models, as well as data collection
practices, which should be all considered when comparing literature values. Incorporation of epidemio-
logical theory into quantitative genetics provides an opportunity to better determine the level of genetic
variation in disease traits, as it accounts for disease transmission among contemporaries. Colostrum
intake is a major determinant of whether a calf develops either respiratory disease or diarrhoea.
Colostrum traits have the advantage of being measured and reported on a continuous scale, which
removes the issues classically associated with binary disease traits. Overall, genetic selection for
improved calf health is feasible. However, to ensure the maximum response, first steps by any industry
members should focus efforts on standardising recording practices and encouragement of uploading
information to genetic evaluation centres through herd management software, as high-quality pheno-
types are the backbone of any successful breeding programme.
Ó 2024 The Authors. Published by Elsevier B.V. on behalf of The Animal Consortium. This is an open access
article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

Implications and recording practices. Most importantly, emphasis should be

placed on encouraging consistent accurate data recording going
The dairy industry is trending away from conventional calf- forward. Furthermore, alternative approaches such as incorporat-
rearing practices, meaning alternative methods to control calf dis- ing epidemiological modelling theory with current quantitative
ease are needed. Genetic selection can be a valuable tool, and cur- genetic approaches, and selection on colostrum traits should be
rent literature shows the potential for calf disease genetic considered as potential avenues to further improve calf health.
evaluation. However, large ranges in heritability estimates exist,
mostly due to differences in model approach, population structure
Introduction

⇑ Corresponding author. In recent years, there has been a notable shift towards prioritis-
E-mail address: [email protected] (C.F. Baes). ing animal health, driven by increasing awareness of how diseases

https://doi.org/10.1016/j.animal.2024.101141
1751-7311/Ó 2024 The Authors. Published by Elsevier B.V. on behalf of The Animal Consortium.
This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
C. Lynch, E.M. Leishman, F. Miglior et al. Animal 18 (2024) 101141

can negatively impact dairy cattle performance. Additionally, there vide a practical framework for characterising respiratory disease, it
is a growing interest of the public in agricultural production can be split into three distinct groups, as described by Callan and
(Svensson and Hultgren, 2008; Alonso et al., 2020). For example, Garry (2002). Firstly, respiratory disease encompasses conditions
in Canada, national genetic evaluations for dairy cattle now contain such as shipping fever pneumonia and enzootic calf pneumonia
various health-related traits, including mastitis resistance, ketosis, where bacterial and viral agents attack the respiratory tract of
hoof lesions, displaced abomasum, fertility disorders, and more the animal. Second is acute interstitial pneumonias which refers
(Koeck et al., 2012; Beavers and Van Doormaal, 2013, 2018; to acute inflammatory conditions affecting the interstitial tissue
Jamrozik et al., 2016, 2021; Miglior et al., 2016; Malchiodi et al., of the lungs more selectively. Lastly, metastatic pneumonia refers
2017; Fleming and Van Doormaal, 2020). However, these traits to infections that spread to the lungs from a primary site of infec-
are measured on adult cows; early-life health traits have received tion in another part of the body. This can occur when pathogens
comparatively less attention until recently. Several progressive enter the bloodstream and reach the lungs, leading to secondary
studies have highlighted the potential of genetic selection to pneumonia. As evident, diarrhoea and respiratory disease can be
improve calf health (Henderson et al., 2011; Gonzalez-Peña et al., caused by a multitude of pathogens making control of the diseases
2019, 2020; Haagen et al., 2021; Zhang et al., 2022; van very difficult on farm. This is apparent when considering the inci-
Staaveren et al., 2023; Lynch et al., 2024). Additionally, Zoetis dence rates for both diseases across the literature, with diarrhoea
became the first commercial animal health company to include calf generally reported between 23 and 44%, while respiratory disease
disease traits within a genomic test, as part of CLARIFIDEÒ Plus ranged from 12–22% (Windeyer et al., 2014; Urie et al., 2018;
(Zoetis, 2022). Although these efforts are promising, to our knowl- USDA, 2018).
edge, there is no national genetic evaluation for calf disease traits Disease expression is further complicated when considering the
to date. Conventional methods to control calf disease have centred components that control it: the interaction of the pathogen causing
on the early removal of the calf from the dam and rearing calves in the disease, the host’s natural resistance to the pathogen, and the
single pens (Flower and Weary, 2003; Beaver et al., 2019), with environment in which both live (Leblanc et al., 2006). Management
consequently limited interaction with conspecifics and reduced practices can help alleviate pathogen burden in the environment,
disease transmission (Ventura et al., 2013; Pempek et al., 2017). including management of the calving pen, bedding material, hous-
Despite reduced disease transmission, the dairy industry may be ing dynamics, and feeding (Renaud et al., 2018; Karle et al., 2019).
encouraged to shift back to group housing and increased cow- Efforts to enhance host resistance have primarily revolved around
calf time for several reasons. These reasons include poor public vaccine programmes and prophylactic antibiotic treatment
opinion of calf social isolation and early removal (Hötzel et al., (Woolums, 2021; Uyama et al., 2022). Vaccine programmes are
2017; Placzek et al., 2021), the demonstrated benefits of group commonplace in dairy herds, with approximately 75% of herds vac-
housing for social support (Duve and Jensen, 2012; Costa et al., cinating against bovine viral diarrhoea in the USA (APHIS, 2008).
2016), and improved feeding behaviour of group-housed calves Roughly 25% of calves receive antimicrobial treatment, primarily
(De Paula Vieira et al., 2012; Costa et al., 2015; Jensen et al., due to diarrhoea or respiratory disease cases. Among calves diag-
2015). Additionally, the other conventional method of treating calf nosed with diarrhoea or respiratory disease, 69 and 88% receive
diseases with antibiotics is not necessarily desirable long term due antimicrobial treatment, respectively (Urie et al., 2018). However,
to concerns with antibiotic use and antimicrobial resistance, which growing global concern around the emergence of antibiotic-
further underlines the need for other strategies to improve calf resistant bacteria and vaccine-resistant viruses, coupled with their
health. Therefore, calf health will become an even bigger area of often-limited effectiveness at preventing disease transmission,
interest and genetic selection may become a valuable tool to diminishes the attractiveness of these interventions (Gibson and
reduce disease incidence in dairy calves. Bishop, 2005; De Oliveira et al., 2020).
Before discussing potential avenues for enhancing calf health In addition to respiratory disease and diarrhoea being highly
through genetic selection, the fundamental elements of calf health, prevalent and difficult to control, previous work has also estab-
diseases, and effects on performance should be discussed. Two lished that calf disease negatively impacts production efficiency.
major calf disease classes are gastrointestinal disease (henceforth Nor et al. (2013) reported a 6% increase in rearing costs in heifers
referred to as diarrhoea) and respiratory disease. Diarrhoea is char- exhibiting disease during calfhood. Sischo et al. (1990) found that
acterised by persistent loose stool lasting multiple days, accompa- calf disease accounted for 4% of total lifetime costs. Estimates for
nied by signs such as decreased appetite, lethargy, dehydration, the cost per case of diarrhoea ranged between 73 and 128 USD
and fever (Renaud et al., 2020; Springer and Yost, 2022). This dis- (Pro Earth Animal Health, 2017; Roche et al., 2020), while the cost
ease primarily affects calves within the 1st 2–3 weeks of life, with per incident of respiratory disease case ranged between 42 and 282
the timing of infection often associated with specific pathogens USD (Dubrovsky et al., 2020; Overton, 2020). Cost variability is due
(Blanchard, 2012). Common pathogens implicated in diarrhoea to several factors, including the severity of the disease, the age of
can be viral (e.g., bovine rotavirus, bovine coronavirus, bovine viral the calf, and management practices (e.g., disease intervention
diarrhoea virus), parasitic (e.g., Cryptosporidium parvum, Eimeria strategies). The associated costs of these diseases not only arise
spp.) and bacterial (e.g., Escherichia coli K99, Salmonella spp., from medical intervention and labour costs but also from effects
Clostridium perfringens) (Foster and Smith, 2009; Cho and Yoon, on production such as average daily gain and future milk potential.
2014). Escherichia coli K99 generally affects calves within the 1st In a meta-analysis of 27 studies, Buczinski et al. (2021) reported
3 or 4 days after birth, bovine rotavirus is most infectious between that calfhood respiratory disease reduced heifer average daily gain
4 and 14 days old, Cryptosporidium parvum and bovine coronavirus by 0.067 kg/d. For diarrhoea, the literature is less definitive, with
are commonly observed between the first and 3rd week of life, and studies reporting a decrease of up to 0.03 kg/d (Donovan et al.,
Salmonella can affect a calf at any point in its early life (McGuirk, 1998; Stanton, 2011), while others found no significant effect
2008; Foster and Smith, 2009; Brunauer et al., 2021). (Virtala et al., 1996). Concerning production, cows with a single
Respiratory disease encompasses infections that affect both the case of diarrhoea when they were calves produced 344 kg less
upper and lower respiratory tracts and can be caused by both energy-corrected milk than healthy animals during the first lacta-
viruses and bacteria (Gorden and Plummer, 2010). Respiratory dis- tion (Svensson and Hultgren, 2008). Similarly, heifers experiencing
ease commonly occurs before the age of 6 months, especially when respiratory disease produced 121.2 kg less milk during their first
maternal antibodies begin to diminish, combined with environ- lactation compared to healthy animals (Buczinski et al., 2021).
mental stressors (Donovan et al., 1998; Taylor et al., 2010). To pro- These findings show the influence of calf illnesses on animals mak-
2
C. Lynch, E.M. Leishman, F. Miglior et al. Animal 18 (2024) 101141

ing the transition from calfhood to the milking herd and help using the method described in the appendix of Dempster and
explain why respiratory disease and diarrhoea are the primary rea- Lerner (1950) by Alan Robertson:
sons behind involuntary culling and mortality until first calving 2
(Waltner-Toews et al., 1986; Wathes et al., 2008; Zhang et al., 2 hl z2
ho ¼
2019b). Preweaning mortality rates in dairy calves range from 5– pð1  pÞ
11% (Murray, 2011; Urie et al., 2018; Winder et al., 2018; Zhang 2 2
et al., 2019b), with diarrhoea (53–56%) and respiratory disease where ho is heritability on the observed scale, hl is heritability on
(21–23%) accounting for roughly 75% of preweaned calf mortality the underlying liability scale, p is the proportion of affected individ-
(NAHMS, 2007; Murray, 2011). For the postweaning period, mor- uals in the population of interest, z is the height of the standard nor-
tality is between 2 and 7%, of which respiratory disease accounts mal curve at the truncation threshold for the corresponding value of
for roughly 50% of cases (NAHMS, 1996; Winder et al., 2018; p. Theoretically, the threshold model is better for disease traits
Santman-Berends et al., 2019; Zhang et al., 2019b; Peel, 2020). because it respects the discrete nature of the trait by fitting a
Clearly, calf disease can have a substantial impact on farm pro- non-Gaussian distribution (Roff, 1996). However, with large num-
ductivity, and that societal concerns regarding current rearing bers of records, model fit and animal ranking do not differ signifi-
practices of dairy calves need to be addressed. Genetic selection cantly between linear and threshold models. This lack of
may be a tool to address these issues long term and needs to be difference means both models are viable options for the genetic
further explored. The objective of this review is to examine the cur- evaluation of disease traits (Jamrozik et al., 2005; Koeck et al.,
rent literature on genetic selection for enhancing calf health and to 2012). Across the literature, heritability estimates on the observed
discuss the key opportunities and challenges that are likely to arise scale for diarrhoea ranged from 0.03 to 0.20, while those for respi-
in implementing routine genetic selection. This review compares ratory disease estimated ranged from 0.02 to 0.24 (Table 1). These
current genetic estimates, exploring the impact of recording prac- large ranges can be attributed to differences in disease prevalence,
tices and calf-specific information on model estimates. Further- population structure, data editing and model approach, and/or data
more, alternative approaches to improve calf health via collection practices. This section will highlight how each of these
epidemiological modelling approaches, and the potential of indi- points can lead to differences in reported values. Furthermore, the
rectly selecting for improved calf health through improving colos- value of incorporating genomic information into genetic evalua-
trum quality are discussed. tions of calf disease traits is discussed.

Effect of prevalence and population structure

Comparing current genetic approaches
The linear model, which estimates parameters on the observed
Current genetic parameters in the literature for diarrhoea and scale, does not account for prevalence. Heritability estimates when
respiratory disease are summarised in Table 1. Studies that esti- using a linear model are maximised when prevalence nears 50%,
mated parameters using a threshold model had their values con- and approach 0 when prevalence nears 0 or 100%. This is because
verted to the observed scale. This transformation was completed phenotypic variance of binary disease traits is determined by the

Table 1
Summary of genetic parameters in the literature for diarrhoea and respiratory disease in dairy calves.

Paper Country Breed Diagnosis Traits Sample size Prevalence Minimum Model 2
hl
2
ho
(# herds) disease
incidence
Lynch et al., 2024 Canada Holstein Producer RESP 158 273 (288) 19% 1% Linear NA 0.03
Threshold 0.05 0.02
102 438 (180) 26% 5% Linear NA 0.04
Threshold 0.06 0.03
DIAR 65 642 (122) 19% 1% Linear NA 0.04
Threshold 0.07 0.03
39 965 (61) 28% 5% Linear NA 0.06
Threshold 0.10 0.05
Zhang et al., 2022 China Holstein Veterinarian RESP 184 563 (31) 4% NA Linear NA 0.05
Threshold 0.08 0.02
DIAR 12% Linear NA 0.06
Threshold 0.10 0.04
Yin et al., 2022 Germany Holstein Producer & Veterinarian RESP 112 563 (53) 28% NA Threshold 0.10 0.06
DIAR 176 904 (53) 22% 0.17 0.09
Haagen et al., 2021 USA Holstein Producer RESP 10 527 (16) 12% 1% Threshold 0.10 0.04
DIAR 11 603 (16) 44% 0.08 0.05
Quick et al., 2020 USA Holstein Veterinarian RESP 1 017 (6) 19% NA Linear NA 0.24
Johnston et al., 2020 Ireland Mixed Producer RESP 727 (32) 5% NA Linear NA 0.09
DIAR 24% NA 0.20
Gonzalez-Peña et al., 2020 USA Jersey Producer RESP 276 134 (90) 14% 0.5% Threshold 0.06 0.02
DIAR 186 505 (90) 37% 0.08 0.05
Gonzalez-Peña et al., 2019 USA Holstein Producer RESP 1 331 626 (188) 21% 0.5% Threshold 0.04 0.02
DIAR 741 484 (118) 26% 0.05 0.03
Mahmoud et al., 2017 Germany Holstein Producer & Veterinarian RESP 31 396 (43) 28% NA Threshold 0.07 0.04
DIAR 21% 0.06 0.03
Neibergs et al., 2014 USA Holstein Laboratory RESP 2 763 (NA) 50% NA Linear NA 0.13
McCorquodale et al., 2013 Canada Holstein Veterinarian RESP 1 588 (16) 10% NA Linear NA 0.04
Henderson et al., 2011 Canada Holstein Veterinarian RESP 7 372 (36) 38% NA Linear NA 0.10
2 2
RESP = Respiratory disease; DIAR = Diarrhoea; hl = Heritability on liability scale; ho = Heritability on observed scale.

3
C. Lynch, E.M. Leishman, F. Miglior et al. Animal 18 (2024) 101141

prevalence of the disease. If prevalence is close to 0 or 100%, there for recording calf disease traits. The first step when adding a new
is limited phenotypic variance, while as prevalence reaches 50% it phenotype of interest to genetic evaluations is to define the trait.
is maximised, better-enabling prediction of genetic differences This is a challenge when it comes to both diarrhoea and respiratory
between animals. Furthermore, at these extremes in prevalence, disease, as laboratory testing is required for accurate etiologic
genetic variation is largely due to non-additive effects, namely epi- diagnosis (Cho and Yoon, 2014). Since this type of analysis is not
static effects (Dempster and Lerner, 1950). The threshold model feasible for most farmers, visual appraisal and clinical examination
estimates parameters on the liability scale, which accounts for by either farmers or veterinarians is the common practice
prevalence in the population. This explains why liability scale her- (McGuirk and Peek, 2014). Common signs of diarrhoea include
itability estimates are larger. Observed scale and linear scale esti- watery faeces and lethargy (Renaud et al., 2020), while respiratory
mates will become closer as disease prevalence nears 50% in a disease signs include coughing, fever, nasal discharge, and changes
study population. For example, Zhang et al. (2022) had a preva- in respiration rate (McGuirk and Peek, 2014). Given the subjective
lence of 4% for respiratory disease, and in this study, the difference nature of visual appraisal, variation in definition can exist, leading
between the observed scale heritability estimate and the liability to many cases being misdiagnosed. Producers tend to diagnose
scale heritability estimate was 4 times smaller (0.02 vs 0.08) cases with relatively low sensitivity (more likely to miss identify-
(Table 1). In contrast, Haagen et al. (2021) had a diarrhoea preva- ing sick animals), but higher specificity (more likely to correctly
lence of 44%, and the difference between the observed and liability identify healthy animals) (Sivula et al., 1996). Knauer et al.
scale estimates was 1.6 times smaller (0.05 vs 0.08). Lynch et al. (2017) reported that producers could identify the health status of
(2024) further confirmed the impact of prevalence on observed group-housed calves with a sensitivity of 26% and a specificity of
scale estimates when comparing different data inclusion criteria 97%. Low sensitivity leads to an under-representation of disease
with respect to minimum herd-year disease incidence (1 vs 5%), prevalence due to a higher amount of false negatives. Therefore,
which in turn impacted the prevalence across analyses. From to maximise selection potential, methods to increase recording
Table 1, we can see that the magnitude of the difference between consistency need to be considered. Possible avenues to help
observed and liability scale estimates is less when the analysis improve the consistency of recording are scoring systems, which
had a disease prevalence closer to 0.5. Together, this shows the help remove the subjectivity of the tester, to provide a more objec-
impact that prevalence has on the observed scale estimates, and tive evaluation, and to allow for greater consistency across testers
therefore, should always be considered when comparing heritabil- (Hayes et al., 2010). Probo and Veronesi (2022) and Wilson et al.
ity estimates across studies. (2023) have reviewed current scoring systems. Examples for respi-
Population structure is another factor that can impact the size ratory disease include the Wisconsin scoring system (WIS score)
of the heritability estimates. One issue with calf disease traits is and the California scoring system (CAL score) (McGuirk, 2008;
that accurate recording of information can be difficult, and as a Love et al., 2014). Buczinski et al. (2015) reported a sensitivity
result, parameters are often estimated on relatively small groups and specificity of 62.4 and 74.1% for the WIS score, respectively.
of animals (<3,000) on limited herds (McCorquodale et al., 2013; For the CAL score, sensitivity and specificity were reported to be
Neibergs et al., 2014; Johnston et al., 2020; Quick et al., 2020). 89.4 and 90.8%, respectively (Love et al., 2014). The Kappa coeffi-
The issue with small datasets is that sires may be poorly dis- cient of agreement between the WIS score and CAL score was esti-
tributed across herds, potentially causing a confounded effect mated to be 0.85, indicating excellent agreement between
between sires’ genetic effects and herds’ environmental effect. If methods (Aly et al., 2014). For diarrhoea, most methods centre
a sire only has records from a single herd, it is hard to untangle around faecal consistency scoring (Lewis and Heaton, 1997;
the environmental influence versus the sire’s influence on the phe- Lesmeister and Heinrichs, 2004; Renaud et al., 2020). Faecal consis-
notype, often resulting in inflated estimates either positively or tency has been reported to be used currently by dairy farmers as a
negatively. As additive variance is directly related to the variance decision tool regarding treatment for diarrhoea (McGuirk, 2008).
of the estimated breeding values, this can lead to an overestima- Unfortunately, no studies relating to the sensitivity or specificity
tion of heritability within a population. This effect on heritability of faecal consistency scoring were found in the literature by the
estimates was shown by Neibergs et al. (2014) who analysed two authors, which would be of great value to the industry. Overall,
datasets from California and New Mexico, using exclusively geno- these methods help improve the diagnosis of disease and provide
mic relationships. The heritability estimate for respiratory disease an opportunity to increase the consistency of recording practices.
when locations were analysed separately was 0.21 for both loca- A concerted effort from industry to encourage the use of these
tions. When the locations were analysed together, the heritability methods will further maximise the potential of genetic selection
estimate dropped to 0.13. Both estimates are high relative to other for calf disease traits.
studies with much larger datasets, which may be partially linked to Apart from accurate recording of the disease, collection of speci-
the aforementioned effect of sire distribution. However, it is essen- fic calf and disease information should also be encouraged. Specific
tial to acknowledge that other factors, such as genotype- calf information includes colostrum (quality, quantity, and timing
environment interaction and degree of genetic connectedness of administration), calving difficulties, and calf vitality scoring.
across contemporary groups within the population, may also con- Colostrum intake and successful passive transfer is a crucial deter-
tribute to variations in heritability estimates (Roso et al., 2004; minant as to whether a calf exhibits morbidity and will be dis-
Zhang et al., 2019a). Overall, care needs to be given when inter- cussed in detail later in this review (Mee, 2008; Furman-Fratczak
preting heritability estimates on small samples of animals, as it et al., 2011). In a meta-analysis, (Raboisson et al., 2016) reported
is possible that estimated effects may be biased. that the adjusted risk in calves with failure of passive transport
(FPT) for respiratory disease and diarrhoea were 1.75 and 1.51,
Recording practices respectively. Lombard et al. (2007) reported that calves born to
dams having severe dystocia had greater odds of treatment for res-
The success of long-term genetic selection programmes is dic- piratory disease (1.7) and digestive disease (i.e., diarrhoea) (1.3).
tated by the recording of accurate and high-quality phenotypes Calf vitality scoring considers characteristics of newborn calves
(Coffey, 2020). As mentioned, disease prevalence greatly influences that relate to suckling reflex, standing, time to lift head, and colos-
heritability on the observed binary scale, influencing the potential trum intake (Barrier et al., 2012). Several methods relating to scor-
rate of genetic change. Therefore, efforts should be made to help ing calf vitality have been proposed (Mee, 2008; Murray-Kerr et al.,
develop a cost-effective data pipeline with standardised protocols 2018; Probo and Veronesi, 2022). However, many of these methods
4
C. Lynch, E.M. Leishman, F. Miglior et al. Animal 18 (2024) 101141

have failed to be adopted on a commercial level, likely due to the further calf information, it is possible to greatly improve our heri-
complexity of the scores and missing recommendations for inter- tability estimates and in turn our potential response to selection.
vention (Homerosky et al., 2017; Probo and Veronesi, 2022). Fur- It must be noted that Henderson et al. (2011) estimated heri-
thermore, conflicting relationships between calf morbidity and tability of respiratory disease as part of a multiple trait analysis
calf vigour have been reported (Sorge et al., 2009; Tora et al., with umbilical disease and bloat, and therefore may benefit from
2021; Alemu et al., 2022). From a genetic selection perspective, calf other trait information within their model. The benefits of multiple
vitality scoring at this point will not benefit current approaches. trait analysis for respiratory disease and diarrhoea together have
However, with the development of better indicators of calf vitality, been reported, with genetic correlations between the traits ranging
it may be a resource to consider in the future. from 0.15 to 0.62 (Mahmoud et al., 2017; Gonzalez-Peña et al.,
Specific disease information includes the duration of the dis- 2019; Haagen et al., 2021; Zhang et al., 2022; Lynch et al., 2024).
ease, to enable a classification of the severity of the disease. This As previously mentioned, these large ranges may be a result of dif-
will allow for clearer distinctions between the phenotypes and ferences in prevalence across traits and population structure
may help improve model performance (Lynch et al., 2024). within studies. Of these studies, only Lynch et al. (2024) compared
Throughout this section, we have identified numerous sources of heritability estimates between univariate and multivariate analy-
information that may help improve our potential for genetic selec- sis, in which they reported increases in estimated heritability for
tion of calf health. It must be noted that it is likely unfeasible to respiratory disease when diarrhoea information was included in
expect producers to collect all information noted above, so a con- the model. Biologically, the positive genetic correlation between
certed effort from industry is required to identify which informa- respiratory disease and diarrhoea is logical. Dairy calves experienc-
tion they feel producers can collect routinely and effectively. ing a disease are more likely to suffer a second illness later in life
(McCorquodale et al., 2013). Diarrhoea can lead to dehydration,
anorexia, and impaired immune function (Schinwald et al., 2022),
Data editing and modelling approach which are all risk factors for developing the respiratory disease
(Gorden and Plummer, 2010). Together, these points suggest that
Across the literature, there are differences in model approaches respiratory disease and diarrhoea should be analysed together in
due to factors such as population structure and recording practices. a multiple-trait analysis, to maximise selection potential.
To highlight the impact of these differences when comparing mod-
elling approaches, we can look at the studies by Gonzalez-Peña Genomic information
et al. (2019) and Henderson et al. (2011). For respiratory disease,
these studies had the largest difference in heritability estimates Genomic information has been incorporated into several stud-
(2 vs 10%) (Table 1) when removing studies with small datasets ies of calf health throughout the literature (Neibergs et al., 2014;
(<3 000 records). This discrepancy may be because two very differ- Mahmoud et al., 2017; Gonzalez-Peña et al., 2019, 2020;
ent samples of animals were analysed, as we know population Johnston et al., 2020; Quick et al., 2020; Haagen et al., 2021; Yin
structure and the relatedness of animals can have a large impact et al., 2022). For variance component estimation, the inclusion of
on variance components. Further reasons for this difference could genomic information was found to show little difference compared
be due to collection practices, incidence thresholds, and model fac- to conventional pedigree-based methods (Gaddis et al., 2014;
tors. Gonzalez-Peña et al. (2019) analysed records from 183 herds Gonzalez-Peña et al., 2019). The value of genomic information is
compared to Henderson et al. (2011) who had phenotypes col- by increasing the accuracy of selection, particularly in low heri-
lected by veterinarians at a single facility that reared calves from tability traits, such as calf disease (VanRaden et al., 2009). Conven-
36 farms. Therefore, Henderson et al. (2011) had the advantage tionally, for low heritability traits, sires required many phenotypes
of trained professionals for diagnosing respiratory disease, and to achieve a high accuracy estimated breeding value. However,
there was a high level of consistency in diagnosis since all animals through genomics, high accuracies can be achieved when no phe-
were reared in one facility. Gonzalez-Peña et al. (2019) likely had notypic information is available on an animal, provided the refer-
variation in definition and recording practices across herds within ence population is representative of the current population of
their analysis. Gonzalez-Peña et al. (2019) set a minimum herd- animals (García-Ruiz et al., 2016; Wiggans et al., 2017). Young sire
year disease incidence of 0.5%, while it was not specified for reliabilities have nearly doubled since the introduction of genomic
Henderson et al. (2011) we assume this was because they were selection across all traits (Fleming and Van Doormaal, 2022). Selec-
all reared at the same facility. The impact of including herds with tion decisions can therefore be made earlier in life, reducing the
very low incidence revolves around the impact of prevalence on generation interval, and leading to an increased response to selec-
model estimates. Furthermore, though it is not stated how many tion (Schefers and Weigel, 2012). As a result of genomic selection,
herds had very low incidence, it would be very rare for a herd to the rate of genetic gain has been doubled. For example, the net
have an incidence around 1%, with between 5 and 15% reported merit index has increased at 85 USD per year since 2010, compared
as the gold standard target for both respiratory disease and diar- to 40 USD in the previous 5 years (Wiggans and Carrillo, 2022). In
rhoea (Dairy Calf and Heifer Association, 2019; RSPCA, 2021). In Canada, the Pro$ index increased at 102 CAD per year prior to
most cases, this would be due to under-recording of cases, due to genomics, and 246 CAD per year postgenomics (Fleming and Van
reasons highlighted in the previous section. Under recording may Doormaal, 2022). Genomics will play a large role in any future
impact model performance as some sick animals that were not genetic evaluation. Therefore, studies should incorporate genomic
recorded will be assumed healthy. If we compare the model factors information if it is available when analysing calf disease traits.
between papers, the major difference is that Henderson et al.
(2011) incorporated specific calf information, including calf weight Limitations of current approaches
upon arrival at the rearing facility and serum total protein informa-
tion (as an indicator for successful passive transfer). This added Throughout this section, we have highlighted current practices
information further accounts for environmental variation associ- that are being suggested and implemented for improving dairy calf
ated with calf diseases, increasing the heritability estimate of the health. However, there are several limitations to these current
trait. The aim here is not to suggest superiority in approach from approaches such as: (1) the effect of prevalence and population
either study but to highlight the possibilities with these traits structure on estimates, leading to inflated genetic parameter val-
and to show that with excellent data collection and collection of ues that likely are not representative of the population, (2) the lack
5
C. Lynch, E.M. Leishman, F. Miglior et al. Animal 18 (2024) 101141

of calf information, such as colostrum intake and dystocia, which a disease outbreak can happen. If it is less than one, the epidemic
are known to play a pivotal role in calf disease expression, and will die out, but if it exceeds one, outbreaks may occur
(3) limited recording practices to ensure consistent accurate infor- (Diekmann et al., 1990; Anche et al., 2014; Tsairidou et al., 2019).
mation is being recorded across herds. Even though there are The basic reproductive ratio is controlled by the transmission rate
opportunities to improve each of these challenges, alternative of the disease and the resulting recovery rate from infection. Across
strategies to improve calf health exist, which are discussed in the the literature, most studies have concentrated on the genetics of
following section. the transmission rate, which is determined by two host traits,
the host’s susceptibility to contracting a disease, and a host’s infec-
tivity to transmit the disease (Anche et al., 2014; Lipschutz-Powell
Alternative approaches
et al., 2014; Biemans et al., 2017; Tsairidou et al., 2019). From an
epidemiology perspective, both susceptibility and infectivity can
We have outlined the current genetic approaches to improve
be seen as indirect genetic effects (IGEs), which are when the geno-
calf health. However, as mentioned, there are several limitations
type of one individual affects the phenotype of a second individual
with current strategies. A number of intriguing avenues of research
(Griffing, 1967; Muir, 2005; Bijma, 2014). These IGE can drastically
have emerged to investigate novel or alternative strategies to
affect the rate and direction of response to selection, which has
improve calf health. We have identified two main areas of interest:
been shown both theoretically and experimentally (Griffing,
(1) epidemiological approaches and (2) investigating the role of
1976; Lipschutz-Powell et al., 2012; Anche et al., 2014). One reason
colostrum, which are discussed in the following sections.
for this is that greater genetic variation may exist in infectivity
compared to susceptibility, enabling faster rates of genetic change.
Epidemiological approach Across generations, natural selection has reduced genetic variation
in susceptibility, as it is linked to an individual’s fitness (Lipschutz-
The ultimate goal of disease control is eradication of the disease Powell and Woolliams, 2012). However, infectivity is not linked to
whereby all animals remain healthy. However, this is theoretically an individual’s own fitness, and therefore, genetic variation would
impossible to achieve from a classical quantitative genetics per- not have been impacted by natural selection (Boddicker et al.,
spective, as additive genetic variance on the observed binary scale 2012).
will approach 0 as prevalence approaches 0 (Dempster and Lerner, Epidemiological models unravel the genetic heterogeneity in
1950). As mentioned previously, this is because when prevalence is both susceptibility and infectivity traits, and respect the impact
close to 0, there is limited phenotypic variance, reducing your abil- each animal has on its contemporaries compared to conventional
ity to differentiate between animals genetically (Hulst et al., 2021). quantitative genetic approaches (Nath et al., 2008; Doeschl-
Furthermore, current methods focus on the susceptibility of indi- Wilson et al., 2011). From a calf health perspective, there is a shift
viduals to disease, disregard information from group members towards group housing of young livestock which increases the
(Lipschutz-Powell et al., 2014), and assume that the observed phe- interaction of animals from a young age (Costa et al., 2015;
notypic differences represent differences in host resistance to a Jensen et al., 2015; Placzek et al., 2021). It would be possible to
given pathogen (Lipschutz-Powell et al., 2013). This approach is apply epidemiological theory discussed above through the record-
unrealistic though, as it assumes that exposure to a pathogen is ing of animal groups/pens to determine which animals were
constant over time, equal among individuals, and purely due to exposed to each other at each point of calfhood. This of course
the environment (Lipschutz-Powell et al., 2013). For this reason, becomes difficult from a management perspective to continuously
it is likely that conventional genetic methods are capturing only track, especially on large enterprises. However, with continuous
a fraction of genetic variation of disease data (Lipschutz-Powell incorporation of precision livestock technology, such as sensor
et al., 2012; Bishop and Woolliams, 2014). data, it may be feasible to automatically track calves via ear tags,
The incorporation of epidemiological theory into quantitative to record calf location at any point in time, and is already commer-
genetics provides an opportunity to better determine the level of cially available for adult cows (Wolfger et al., 2017; Costa et al.,
genetic variation in disease traits (Bishop and Stear, 1997; Bijma 2021a).
et al., 2022). This stems from the ability to allow for the positive
feed-back dynamics of disease transmission to be accounted for Colostrum
(Anche et al., 2014; Bijma et al., 2022). The basis of this positive
feed-back dynamic is as follows; if an animal is immune to a One of the major determinants for whether a calf develops
pathogen, this has a positive effect on animals around them, as either respiratory disease or diarrhoea is the quantity and quality
they will not spread the pathogen. If many animals within a group of colostrum that they received and when they receive it. There-
are immune, this positive feedback effect will be compounded, to fore, a possible proactive measure to help reduce calf disease
the point where the pathogen can no longer spread, and thus die would be genetically selecting for traits related to colostrum qual-
out completely. On the other hand, if an animal is very susceptible ity of the dam and antibody absorption of the calf. The importance
to a disease, they will become sick and aid in the spread of disease. of colostrum feeding in calf rearing has been well-documented, as
This increases the likelihood of animals around them becoming it profoundly influences health and long-term performance (Khan
sick, potentially resulting in an outbreak within a group of animals et al., 2011; Ballou, 2012; Hammon et al., 2020). Colostrum is the
(Bijma et al., 2022). An example of this positive feedback effect is first milk that a cow releases postcalving, and is rich in antibodies,
seen with herd immunity via vaccination protocols, whereby com- also known as immunoglobulins (Ig), and other nutrients that pro-
plete eradication of a disease can be achieved without the full pop- vide the calf protection from disease early in life (McGrath et al.,
ulation being vaccinated, as the pathogen is unable to spread (Fine, 2016). The importance of feeding colostrum stems from dairy cows
1993). not having an epitheliochorial placenta, meaning passive immu-
The key parameter in epidemiology to access this positive feed- nity cannot be transferred to the neonate during gestation
back mechanism and the impact of disease interventions on epi- (Robbers et al., 2021). Therefore, calves solely rely on the transfer
demic risk is the basic reproductive ratio, which represents the of immune constituents through initial feedings from the dam
average number of secondary cases caused by an infected individ- via colostrum. Within colostrum, there are three major Ig, IgG
ual in a susceptible population (Diekmann et al., 1990). The basic (80–90%) is considered the most important, while IgM and IgA
reproductive ratio has a threshold value of one, which dictates if make up about 7 and 5%, respectfully (Stilwell and Carvalho,
6
C. Lynch, E.M. Leishman, F. Miglior et al. Animal 18 (2024) 101141

2011; Godden et al., 2019). Colostrum quality varies among ani- IgG concentration, total solids (Brix %), serum total protein, serum
mals (Gilbert et al., 1988) and is determined by numerous factors, IgG, and numerous associated natural antibodies. Heritability esti-
including volume, collection time, Ig concentration, and bacterial mates for colostrum yield range from 0.04 to 0.21 (Conneely et al.,
levels (McGuirk and Collins, 2004; Godden, 2008). Timing of feed- 2013; Soufleri et al., 2019), IgG concentration ranged from 0.10 to
ing is also important. To ensure successful passive transfer of Ig, 0.40 (Gilbert et al., 1988; Conneely et al., 2013; Cockrum et al.,
the calf must consume colostrum shortly after birth, ideally within 2016; Martin et al., 2021; Cordero-Solorzano et al., 2022), and total
2–3 h (Weaver et al., 2000; Godden, 2008). If colostrum feeding is solids (Brix %) ranged from 0.27 to 0.41 (Soufleri et al., 2019;
delayed or is of low quality, FPT can occur (Raboisson et al., 2016). Cordero-Solorzano et al., 2022). For calf absorption ability, the her-
FPT occurs when the calf serum IgG concentration falls below itability estimate for serum total protein was 0.07 (Cordero-
10 mg/mL (Pritchett et al., 1991; Faber et al., 2005; Beam et al., Solorzano et al., 2022), while serum IgG estimates ranged from
2009), and is associated with increased calf mortality and morbid- 0.18 to 0.56 (Gilbert et al., 1988; Burton et al., 1989; Martin
ity rates (Mee, 2008; Furman-Fratczak et al., 2011). FPT rates vary et al., 2021; Cordero-Solorzano et al., 2022). The large range in esti-
across the literature, falling somewhere between 19 and 40% mates could be due to many factors including differences in popu-
(Beam et al., 2009; Raboisson et al., 2016; Abuelo et al., 2019). lation structure, colostrum and serum measurement methods, and
Together, this highlights the importance of colostrum in calf dis- phenotyping strategies. However, it is clear that there are much
ease prevention. Furthermore, it is common practice in many herds higher heritability estimates for colostrum traits compared to
to pool colostrum to help reduce labour requirements (King et al., health traits, which are typically below 0.1. Colostrum traits have
2020), and therefore, improving colostrum quality at the cow level the advantage of being a continuous variable which removes the
may have greater effects at the herd level. However, the benefits of issues classically associated with binary disease traits. The advan-
pooling colostrum must be weighed against the potential for dis- tage of this is due to the clearer distinction between animals phe-
ease transmission, especially in herds with endemic infectious dis- notypically, which increases the ability to identify genetic
eases such as Johne’s Disease or Bovine Leukaemia Virus (Godden, differences amongst animals. However, in many cases, heritability
2008; Williams et al., 2014). estimates were not different from zero, with many studies report-
To evaluate if a trait is suitable for genetic selection, certain cri- ing high SEs. This is likely due to the limited amount of data within
teria should be considered, as described by Shook (1989). This certain studies. Therefore, further research is still required to bet-
includes that the trait needs to be clearly defined, cost-effective ter determine the genetic influence on colostrum-related traits.
and easy to measure, it must exhibit significant genetic variation The last criterion for genetic selection relates to the economic
and be heritable, and there must be economic value to improving importance of the trait. When discussing colostrum traits, we are
the trait. Colostrum quality can be defined by the concentration mainly concerned with helping to reduce FPT by maximising the
of Ig within colostrum, while calf absorption ability can be defined quality of colostrum and the calf’s ability to absorb the associated
as the level of Ig within calf serum 24 h postfeeding. Colostrum antibodies, so economics should be related to the impact of FPT.
quality and calf absorption ability can be directly determined by Raboisson et al. (2016) reported that the average marginal cost
radial immunodiffusion (RID), which is considered the gold stan- for FPT ranged from €60 to €121 for dairy calves and €80 to €140
dard measurement (Bartens et al., 2016; Ahmann et al., 2021). for beef calves. These values stem from the fact that FPT is a lead-
Though RID is highly accurate and repeatable, it is expensive and ing reason behind preweaning morbidity and in-turn mortality.
time-consuming reducing its merit for a potential measure on a Raboisson et al. (2016) also reported that the adjusted risks of
large-scale genetic programme (Costa et al., 2021b). For colostrum FPT (and 95% confidence intervals) for mortality, respiratory dis-
quality, several other measurement methods have been proposed, ease, diarrhoea and overall morbidity were 2.12 (1.43–3.13), 1.75
including the use of enzyme-linked immunosorbent assay (ELISA) (1.50–2.03), 1.51 (1.05– 2.17) and 1.91 (1.63–2.24), respectively.
(Gelsinger et al., 2015), near-infrared spectroscopy (NIRS) (Rivero Preweaning mortality rates in dairy calves range from 5 to 11%
et al., 2012; Costa et al., 2021b), and Brix refractometer (Quigley (Murray, 2011; Urie et al., 2018; Winder et al., 2018; Zhang
et al., 2013; Bartier et al., 2015). Like RID, ELISA tests are time- et al., 2019b) and in the UK, it was reported that calf mortality
consuming and require lab work. NIRS and Brix provide the most costs roughly 60 million pounds/year (Department for
promise, with both being rapid and cost-effective measures for Environment, 2003). Given this large economic impact, reducing
determining colostrum quality. Genetic and phenotypic correla- FPT is of great interest, showing the value of improving
tions between NIRS and RID were estimated to be 0.85 and 0.77, colostrum-related traits through genetic selection.
respectively (Costa et al., 2021b), while phenotypic correlations Genetic selection for colostrum traits has promise for improving
between Brix and RID have ranged from 0.64 to 0.75 (Bielmann overall calf health. Some challenges going forward will include the
et al., 2010; Quigley et al., 2013; Gelsinger et al., 2015), showing standardisation of recording methods to measure colostrum traits.
good potential as indicator traits of RID values. One challenge with For long-term genetic evaluation, one method should be decided
calf absorption ability is that it requires a blood sample, making upon and encouraged to be used by producers to ensure consis-
recording more difficult and time-consuming, while also being tency of testing practices. Another potential challenge is the issue
quite invasive to the animal. For genetic selection on calf absorp- of hypergammaglobulinemia, which refers to the overproduction
tion ability to be feasible, long-term alternative measures will need of immunoglobulin (Beuvon, 2021) and has been related to dis-
to be identified. One area of interest to predicting calf serum Ig eases in numerous species including humans and dogs (Buadi
concentration is through calf saliva, which is less invasive and et al., 2011; Colopy et al., 2019). Evidence of hypergammaglobu-
easier to record (Johnsen et al., 2019; Silva et al., 2023). Johnsen linemia in dairy cattle were not found by the authors but caution
et al. (2019) reported a strong positive correlation (r = 0.7) between would be suggested when selecting colostrum traits, particularly
saliva IgG and serum IgG. However, all sampled calves in this study related to immunoglobulin production to fully understand its
had serum IgG levels above 10 g/L., meaning it is unknown how impact on the dairy cow.
calves with FPT (i.e., serum IgG < 10 g/L) can be predicted by saliva
IgG. Therefore, further work is required to determine if saliva can
be used as an indicator of FPT. Conclusions
For selection, traits also need to have genetic variance and be
heritable. Though limited, several studies have estimated the her- Management of dairy calf health is likely to become a greater
itabilities of colostrum-related traits, including colostrum yield, area of interest as conventional methods of disease control (e.g.,
7
C. Lynch, E.M. Leishman, F. Miglior et al. Animal 18 (2024) 101141

early removal from the dam, single housing) become less attractive Financial support statement
due to societal pressure and the apparent impact on overall calf
development. One result of this shift will be group housing, which This review was conducted as part of the Resilient Dairy Gen-
will expose young calves to higher levels of pathogen. To combat ome Project (RDGP). We gratefully acknowledge all funding and
the potential impact of this exposure, genetic selection may be a support for the Resilient Dairy Genome Project from the organisa-
valuable asset. Throughout this review, we have detailed current tions listed at http://www.resilientdairy.ca/funders-and-partners/,
genetic practices to help improve calf health and highlighted the as administered by Genome Canada, Genome Alberta, Ontario
areas where we see opportunities and challenges, most notably Genomics, Genome Quebec, and Genome British Columbia. Dr.
with respect to improving data recording practices. Furthermore, Christine Baes also gratefully acknowledges support from the Nat-
we have detailed alternative approaches, such as incorporating ural Sciences and Engineering Research Council of Canada (NSERC)
epidemiological modelling to better understand disease transmis- Canada Research Chair and Discovery programmes. As per the
sion, and reviewed the potential of selection on colostrum traits research agreement, the funders had no role in the design and con-
as a proactive measure to improve calf health. Overall, enhancing duct of the study as well as the decision to publish.
calf health through genetic selection is attainable. However, to
optimise response, initial actions by industry stakeholders should Transparency Declaration
prioritise the standardisation of recording practices and the pro-
motion of data uploads to herd management software, as high- This article is part of a supplement entitled Selected keynote
quality phenotypes are the backbone of any successful breeding lectures of the 74th Annual Meeting of the European Federation
programme. of Animal Science (Lyon, France) supported by the Animal
Consortium.

Ethics approval
References
Not applicable. Abuelo, A., Havrlant, P., Wood, N., Hernandez-Jover, M., 2019. An investigation of
dairy calf management practices, colostrum quality, failure of transfer of
passive immunity, and occurrence of enteropathogens among Australian dairy
farms. Journal of Dairy Science 102, 8352–8366.
Data and model availability statement Ahmann, J., Steinhoff-Wagner, J., Büscher, W., 2021. Determining immunoglobulin
content of bovine colostrum and factors affecting the outcome: a review.
Not applicable. Information can be made available from the Animals 11, 1–29.
Alemu, Y.F., Jemberu, W.T., Mekuriaw, Z., Abdi, R.D., 2022. Incidence and predictors
authors upon request.
of calf morbidity and mortality from birth to 6-months of age in dairy farms of
Northwestern Ethiopia. Frontiers in Veterinary Science 9, 1–12.
Alonso, M.E., González-Montaña, J.R., Lomillos, J.M., 2020. Consumers’ concerns and
Declaration of Generative AI and AI-assisted technologies in the perceptions of farm animal welfare. Animals 10, 1–13.
Aly, S.S., Love, W.J., Williams, D.R., Lehenbauer, T.W., Van Eenennaam, A., Drake, C.,
writing process Kass, P.H., Farver, T.B., 2014. Agreement between bovine respiratory disease
scoring systems for pre-weaned dairy calves. Animal Health Research Reviews
During the preparation of this work the author(s) did not use 15, 148–150.
Anche, M.T., De Jong, M.C.M., Bijma, P., 2014. On the definition and utilization of
any AI and AI-assisted technologies.
heritable variation among hosts in reproduction ratio R 0 for infectious diseases.
Heredity 113, 364–374.
APHIS 2008. Bovine Viral Diarrhea (BVD) Management practices and detection in
Author ORCIDs bulk tank milk in the United States, 2007. Animal & Plant Health Inspection
Service- Veterinary Services. Retrieved on 9 January 2023 from https://www.
aphis.usda.gov/animal_health/nahms/dairy/downloads/dairy07/Dairy07_is_
CL: https://orcid.org/0000-0002-0524-3939. BVD_1.pdf.
EL: https://orcid.org/0000-0003-1739-3915. Ballou, M.A., 2012. Immune responses of Holstein and Jersey calves during the
preweaning and immediate postweaned periods when fed varying planes of
FM: https://orcid.org/0000-0003-2345-8842. milk replacer. Journal of Dairy Science 95, 7319–7330.
DK: https://orcid.org/0000-0001-9606-7602. Barrier, A.C., Ruelle, E., Haskell, M.J., Dwyer, C.M., 2012. Effect of a difficult calving
FS: https://orcid.org/0000-0001-8700-0633. on the vigour of the calf, the onset of maternal behaviour, and some behavioural
indicators of pain in the dam. Preventive Veterinary Medicine 103, 248–256.
CB: https://orcid.org/0000-0001-6614-8890.
Bartens, M.-C., Drillich, M., Rychli, K., Iwersen, M., Arnholdt, T., Meyer, L., Klein-
Jöbstl, D., 2016. Assessment of different methods to estimate bovine colostrum
quality on farm. New Zealand Veterinary Journal 64, 263–267.
CRediT authorship contribution statement Bartier, A.L., Windeyer, M.C., Doepel, L., 2015. Evaluation of on-farm tools for
colostrum quality measurement. Journal of Dairy Science 98, 1878–1884.
Beam, A.L., Lombard, J.E., Kopral, C.A., Garber, L.P., Winter, A.L., Hicks, J.A., Schlater, J.
C. Lynch: Writing – review & editing, Writing – original draft, L., 2009. Prevalence of failure of passive transfer of immunity in newborn heifer
Conceptualization. E.M. Leishman: Writing – review & editing, calves and associated management practices on US dairy operations. Journal of
Dairy Science 92, 3973–3980.
Writing – original draft. F. Miglior: Writing – review & editing.
Beaver, A., Meagher, R.K., von Keyserlingk, M.A.G., Weary, D.M., 2019. Invited
D. Kelton: Writing – review & editing. F.S. Schenkel: Writing – review: a systematic review of the effects of early separation on dairy cow and
review & editing. C.F. Baes: Writing – review & editing, calf health. Journal of Dairy Science 102, 5784–5810.
Beavers L and Van Doormaal B 2013. Official Genetic evaluation for mastitis
Conceptualization.
resistance. Canadian Dairy Network. Retrieved on 9 January 2023 from https://
www.cdn.ca/document.php?id=333.
Beavers L and Van Doormaal B 2018. Genetic selection for improved hoof health is
Declaration of interest now possible ! Canadian Dairy Network. Retrieved on 9 January 2023 from
https://www.cdn.ca/document.php?id=513.
Beuvon, C., 2021. Etiologies of polyclonal hypergammaglobulinemia: a scoping
None. review. European Journal of Internal Medicine 90, 119–121.
Bielmann, V., Gillan, J., Perkins, N.R., Skidmore, A.L., Godden, S., Leslie, K.E., 2010. An
evaluation of brix refractometry instruments for measurement of colostrum
Acknowledgements quality in dairy cattle. Journal of Dairy Science 93, 3713–3721.
Biemans, F., De Jong, M.C.M., Bijma, P., 2017. A model to estimate effects of SNPs on
host susceptibility and infectivity for an endemic infectious disease. Genetics
None. Selection Evolution 49, 1–13.

8
C. Lynch, E.M. Leishman, F. Miglior et al. Animal 18 (2024) 101141

Bijma, P., 2014. The quantitative genetics of indirect genetic effects: a selective Diekmann, O., Heesterbeek, A.P., Metz, J.A.J., 1990. Mathematical biology. Journal of
review of modelling issues. Heredity 112, 61–69. Mathematical Biology 28, 365–382.
Bijma, P., Hulst, A.D., de Jong, M.C.M., 2022. The quantitative genetics of the Doeschl-Wilson, A.B., Davidson, R., Conington, J., Roughsedge, T., Hutchings, M.R.,
prevalence of infectious diseases: hidden genetic variation due to indirect Villanueva, B., 2011. Implications of host genetic variation on the risk and
genetic effects dominates heritable variation and response to selection. prevalence of infectious diseases transmitted through the environment.
Genetics 220, iyab141. Genetics 188, 683–693.
Bishop, S.C., Stear, M.J., 1997. Modelling responses to selection for resistance to Donovan, G.A., Dohoo, I.R., Montgomery, D.M., Bennett, F.L., 1998. Calf and disease
gastro-intestinal parasites in sheep. Animal Science 64, 469–478. factors affecting growth in female Holstein calves in Florida, USA. Preventive
Bishop, S.C., Woolliams, J.A., 2014. Genomics and disease resistance studies in Veterinary Medicine 33, 1–10.
livestock. Livestock Science 166, 190–198. Dubrovsky, S.A., Van Eenennaam, A.L., Aly, S.S., Karle, B.M., Rossitto, P.V., Overton,
Blanchard, P.C., 2012. Diagnostics of Dairy and Beef Cattle Diarrhea. Veterinary M.W., Lehenbauer, T.W., Fadel, J.G., 2020. Preweaning cost of bovine respiratory
Clinics of North America: Food Animal Practice 28, 443–464. disease (BRD) and cost-benefit of implementation of preventative measures in
Boddicker, N., Waide, E.H., Rowland, R.R.R., Lunney, J.K., Garrick, D.J., Reecy, J.M., calves on California dairies: the BRD 10K study. Journal of Dairy Science 103,
Dekkers, J.C.M., 2012. Evidence for a major QTL associated with host response to 1583–1597.
porcine reproductive and respiratory syndrome virus challenge. Journal of Duve, L.R., Jensen, M.B., 2012. Social behavior of young dairy calves housed with
Animal Science 90, 1733–1746. limited or full social contact with a peer. Journal of Dairy Science 95, 5936–
Brunauer, M., Roch, F.F., Conrady, B., 2021. Prevalence of worldwide neonatal calf 5945.
diarrhoea caused by bovine rotavirus in combination with bovine coronavirus, Faber, S.N., Faber, N.E., McCauley, T.C., Ax, R.L., 2005. Case study: effects of
escherichia coli k99 and cryptosporidium spp.: a meta-analysis. Animals 11, 1– colostrum ingestion on lactational performance. The Professional Animal
23. Scientist 21, 420–425.
Buadi, F., Hsing, A.W., Katzmann, J.A., Pfeiffer, R.M., Waxman, A., Yeboah, E.D., Fine, P.E.M., 1993. Herd immunity: history, theory, practice. Epidemiologic Reviews
Biritwum, R.B., Tettey, Y., Adjei, A., Chu, L.W., Demarzo, A., Netto, G.J., 15, 265–302.
Dispenzieri, A., Kyle, R.A., Rajkumar, S.V., Landgren, O., 2011. High prevalence Fleming A and Van Doormaal B 2020. New Genetic Evaluations Coming in
of polyclonal hypergamma-globulinemia in adult males in Ghana, Africa. December 2020. Lactanet. Retrieved on 9 January 2023 from https://www.
American Journal of Hematology 86, 554–558. cdn.ca/document.php?id=544.
Buczinski, S., Ollivett, T., Dendukuri, N., 2015. Bayesian estimation of the accuracy of Fleming A and Van Doormaal B 2022. Lactanet: The Impact of Genomics. Lactanet.
the calf respiratory scoring chart and ultrasonography for the diagnosis of Retrieved on 9 January 2023 from https://lactanet.ca/en/impact-genomics/?
bovine respiratory disease in pre-weaned dairy calves. Preventive Veterinary print=enabled&id=175313.
Medicine 119, 227–231. Flower, F.C., Weary, D.M., 2003. The effects of early seperation on the dairy cow and
Buczinski, S., Achard, D., Timsit, E., Cliniques, S., Vétérinaire, F.D.M., De, M.U., Js, C., calf. Animal Welfare 12, 339–348.
2021. Effects of calfhood respiratory disease on health and performance of dairy Foster, D.M., Smith, G.W., 2009. Pathophysiology of diarrhea in calves. Veterinary
cattle : a systematic review and meta-analysis. Journal of Dairy Science 104, Clinics of North America - Food Animal Practice 25, 13–36.
8214–8227. Furman-Fratczak, K., Rzasa, A., Stefaniak, T., 2011. The influence of colostral
Burton, J.L., Kennedy, B.W., Burnside, E.B., Wilkie, B.N., Burton, J.H., 1989. variation immunoglobulin concentration in heifer calves’ serum on their health and
in serum concentrations of Immunoglobulins G, A, and M in Canadian Holstein- growth. Journal of Dairy Science 94, 5536–5543.
Friesian Calves. Journal of Dairy Science 72, 135–149. Gaddis, K.L.P., Cole, J.B., Clay, J.S., Maltecca, C., 2014. Genomic selection for
Callan, R.J., Garry, F.B., 2002. Biosecurity and bovine respiratory disease. Veterinary producer-recorded health event data in US dairy cattle. Journal of Dairy
Clinics of North America - Food Animal Practice 18, 57–77. Science 97, 3190–3199.
Cho, Y. il, Yoon, K.J., 2014. An overview of calf diarrhea - infectious etiology, García-Ruiz, A., Cole, J.B., VanRaden, P.M., Wiggans, G.R., Ruiz-López, F.J., Van
diagnosis, and intervention. Journal of Veterinary Science 15, 1–17. Tassell, C.P., 2016. Changes in genetic selection differentials and generation
Cockrum, R.R., Speidel, S.E., Salak-Johnson, J.L., Chase, C.C.L., Peel, R.K., Weaber, R.L., intervals in US Holstein dairy cattle as a result of genomic selection.
Loneagan, G.H., Wagner, J.J., Boddhireddy, P., Thomas, M.G., Prayaga, K., Denise, Proceedings of the National Academy of Sciences 113, E3995–E4004.
S., Enns, R.M., 2016. Genetic parameters estimated at receiving for circulating Gelsinger, S.L., Smith, A.M., Jones, C.M., Heinrichs, A.J., 2015. Technical note:
cortisol, immunoglobulin G, interleukin 8, and incidence of bovine respiratory comparison of radial immunodiffusion and ELISA for quantification of bovine
disease in feedlot beef steers. Journal of Animal Science 94, 2770–2778. immunoglobulin G in colostrum and plasma. Journal of Dairy Science 98, 4084–
Coffey, M., 2020. Dairy cows: in the age of the genotype, #phenotypeisking. Animal 4089.
Frontiers 10, 19–22. Gibson, J.P., Bishop, S.C., 2005. Use of molecular markers to enhance resistance of
Colopy, L.J., Rout, E.D., Laura, K.S., Anne, A.S., Moore, A.R., 2019. Immunoglobulin livestock to disease: a global approach Utilising and improving genetic
G4-related disease in a dog. Journal of Veterinary Internal Medicine 33, 2732– resistance to disease. Revue Scientifique et Technique (International Office of
2738. Epizootics) 24, 343–353.
Conneely, M., Berry, D.P., Sayers, R., Murphy, J.P., Lorenz, I., Doherty, M.L., Kennedy, Gilbert, R.P., Gaskins, C.T., Hillers, J.K., Brinks, J.S., Denham, A.H., 1988. Inbreeding
E., 2013. Factors associated with the concentration of immunoglobulin G in the and Immunoglobulin G concentrations in cattle. Journal of Animal Science 66,
colostrum of dairy cows. Animal 7, 1824–1832. 2490–2497.
Cordero-Solorzano, J., de Koning, D.J., Tråvén, M., de Haan, T., Jouffroy, M., Larsson, Godden, S., 2008. Colostrum management for dairy calves. Veterinary Clinics of
A., Myrthe, A., Arts, J.A.J., Parmentier, H.K., Bovenhuis, H., Wensman, J.J., 2022. North America - Food Animal Practice 24, 19–39.
Genetic parameters of colostrum and calf serum antibodies in Swedish dairy Godden, S.M., Lombard, J.E., Woolums, A.R., 2019. Colostrum management for
cattle. Genetics Selection Evolution 54, 1–18. dairy calves. Veterinary Clinics of North America: Food Animal Practice 35, 535–
Costa, J.H.C., Cantor, M.C., Neave, H.W., 2021a. Symposium review: precision 556.
technologies for dairy calves and management applications. Journal of Dairy Gonzalez-Peña, D., Vukasinovic, N., Brooker, J.J., Przybyla, C.A., Denise, S.K., 2019.
Science 104, 1203–1219. Genomic evaluation for calf wellness traits in Holstein cattle. Journal of Dairy
Costa, A., Franzoi, M., Visentin, G., Goi, A., De Marchi, M., Penasa, M., 2021b. The Science 102, 2319–2329.
concentrations of immunoglobulins in bovine colostrum determined by the Gonzalez-Peña, D., Vukasinovic, N., Brooker, J.J., Przybyla, C.A., Baktula, A., Denise, S.
gold standard method are genetically correlated with their near-infrared K., 2020. Genomic evaluation for wellness traits in US Jersey cattle. Journal of
prediction. Genetics Selection Evolution 53, 1–14. Dairy Science 103, 1735–1748.
Costa, J.H.C., Meagher, R.K., von Keyserlingk, M.A.G., Weary, D.M., 2015. Early pair Gorden, P.J., Plummer, P., 2010. Control, management, and prevention of bovine
housing increases solid feed intake and weight gains in dairy calves. Journal of respiratory disease in dairy calves and cows. Veterinary Clinics of North
Dairy Science 98, 6381–6386. America: Food Animal Practice 26, 243–259.
Costa, J.H.C., von Keyserlingk, M.A.G., Weary, D.M., 2016. Invited review: effects of Griffing, B., 1967. Selection in reference to biological groups I. Individual and group
group housing of dairy calves on behavior, cognition, performance, and health. selection applied to populations of unordered groups. Australian Journal of
Journal of Dairy Science 99, 2453–2467. Biological Sciences 20, 127–139.
Dairy Calf & Heifer Association 2019. DCHA Gold Standards. Retrieved on 9 January Griffing, B., 1976. Selection in reference to biological groups V. Analysis of full-sib
2023 from https://calfandheifer.org/wp-content/uploads/2020/09/DCHA_ groups. Genetics 82, 703–722.
GoldStandards_2020_En_WEB-final.pdf. Haagen, I.W., Hardie, L.C., Heins, B.J., Dechow, C.D., 2021. Genetic parameters of calf
De Oliveira, D.M.P., Forde, B.M., Kidd, T.J., Harris, P.N.A., Schembri, M.A., Beatson, S. morbidity and stayability for US organic Holstein calves. Journal of Dairy
A., Paterson, D.L., Walker, M.J., 2020. Antimicrobial resistance in ESKAPE Science 104, 11770–11778.
pathogens. Clinical Microbiology Reviews 33, 1–49. Hammon, H.M., Liermann, W., Frieten, D., Koch, C., 2020. Review: importance of
De Vieira Paula, A., von Keyserlingk, M.A.G., Weary, D.M., 2012. Presence of an older colostrum supply and milk feeding intensity on gastrointestinal and systemic
weaned companion influences feeding behavior and improves performance of development in calves. Animal 14, s133–s143.
dairy calves before and after weaning from milk. Journal of Dairy Science 95, Hayes, G., Mathews, K., Kruth, S., Doig, G., Dewey, C., 2010. Illness severity scores in
3218–3224. veterinary medicine: what can we learn? Journal of Veterinary Internal
Dempster, E., Lerner, I., 1950. Heritability of threshold characters. Genetics 35, 212– Medicine 24, 457–466.
236. Henderson, L., Miglior, F., Sewalem, A., Wormuth, J., Kelton, D., Robinson, A., Leslie,
Department for Environment F and RA (DEFRA) 2003 2003. Improving calf survival. K.E., 2011. Short communication: genetic parameters for measures of calf
Retrieved on 9 January 2023 from http://www.veeru.reading.ac.uk/comp2/ health in a population of Holstein calves in New York State. Journal of Dairy
Comcomp/adobe/calfsurv.pdf. Science 94, 6181–6187.

9
C. Lynch, E.M. Leishman, F. Miglior et al. Animal 18 (2024) 101141

Homerosky, E.R., Caulkett, N.A., Timsit, E., Pajor, E.A., Kastelic, J.P., Windeyer, M.C., and genetic parameters for colostrum and calf serum in Charolais animals.
2017. Clinical indicators of blood gas disturbances, elevated L-lactate Journal of Dairy Science 104, 3240–3249.
concentration and other abnormal blood parameters in newborn beef calves. McCorquodale, C.E., Sewalem, A., Miglior, F., Kelton, D., Robinson, A., Koeck, A.,
The Veterinary Journal 219, 49–57. Leslie, K.E.E., 2013. Short communication: analysis of health and survival in a
Hötzel, M.J., Cardoso, C.S., Roslindo, A., Von, K.M., 2017. Citizens ’ views on the population of Ontario Holstein heifer calves. Journal of Dairy Science 96, 1880–
practices of zero-grazing and cow-calf separation in the dairy industry : does 1885.
providing information increase acceptability ? Journal of Dairy Science 100, McGrath, B.A., Fox, P.F., Mcsweeney, P.L.H., Kelly, A.L., 2016. Composition and
4150–4160. properties of bovine colostrum: a review. Dairy Science & Technology 96, 133–
Hulst, A.D., de Jong, M.C.M., Bijma, P., 2021. Why genetic selection to reduce the 158.
prevalence of infectious diseases is way more promising than currently McGuirk, S.M., 2008. Disease management of dairy calves and heifers. Veterinary
believed. Genetics 217, iyab024. Clinics of North America - Food Animal Practice 24, 139–153.
Jamrozik, J., Fatehi, J., Kistemaker, G.J., Schaeffer, L.R., 2005. Estimates of genetic McGuirk, S.M., Collins, M., 2004. Managing the production, storage, and delivery of
parameters for Canadian Holstein female reproduction traits. Journal of Dairy colostrum. Veterinary Clinics of North America - Food Animal Practice 20, 593–
Science 88, 2199–2208. 603.
Jamrozik, J., Kistemaker, G.J., Van, D.BJ., Fleming, A., Koeck, A., Miglior, F., 2016. McGuirk, S.M., Peek, S.F., 2014. Timely diagnosis of dairy calf respiratory disease
Genetic evaluation for resistance to metabolic diseases in canadian dairy using a standardized scoring system. Animal Health Research Reviews 15, 145–
breeds. Interbull Bulletin 50, 9–16. 147.
Jamrozik, J., Kistemaker, G., Van, D.B., Baes, C., Miglior, F., 2021. Genomic evaluation Mee, J.F., 2008. Newborn dairy calf management. Veterinary Clinics of North
for resistance to fertility disorders in Canadian dairy breeds. Interbull Bulletin America - Food Animal Practice 24, 1–17.
56, 102–110. Miglior F, Finocchiaro R, Malchiodi F, Fleming A, Brito L, Baes CF, Jamrozik J, Martin
Jensen, M.B., Duve, L.R., Weary, D.M., 2015. Pair housing and enhanced milk P, Chesnais J and Schenkel FS 2016. Enhancing the data pipeline for novel traits
allowance increase play behavior and improve performance in dairy calves. in the genomic era: from farms to DHI to evaluation centres. In S. Niklitschek, J.
Journal of Dairy Science 98, 2568–2575. Lama, C. Trejo, B. Wickham, M. Burke, and C. Mosconi, eds. Proceedings of the
Johnsen, J.F., Chincarini, M., Sogstad, Å.M., Sølverød, L., Vatne, M., Mejdell, C.M., 40th ICAR Biennial Session, Puerto Varas, Chili. ICAR, Puerto Varas, Chili, pp. 23–
Hänninen, L., 2019. Salivary IgG levels in neonatal calves and its association to 30.
serum IgG: an observational pilot study. Translational Animal Science 3, 589– Muir, W.M., 2005. Incorporation of competitive effects in forest tree or animal
593. breeding programs. Genetics 170, 1247–1259.
Johnston, D., Mukiibi, R., Waters, S.M., McGee, M., Surlis, C., McClure, J.C., McClure, Murray B 2011. Optimizing Calf Survival at Birth. Ontario Ministry of Agriculture,
M.C., Todd, C.G., Earley, B., 2020. Genome wide association study of passive Food and Rural Affairs. Retrieved on 22 November 2022, from http://www.
immunity and disease traits in beef-suckler and dairy calves on Irish farms. omafra.gov.on.ca/english/livestock/dairy/facts/optbirth.htm.
Scientific Reports 10, 1–10. Murray-Kerr CF, Leslie KE, Godden SM, Knauer WA and Mcguirk SM 2018.
Karle, B.M., Maier, G.U., Love, W.J., Dubrovsky, S.A., Williams, D.R., Anderson, R.J., Development of a newborn calf vigor scoring system. In: Proceedings of the
Van Eenennaam, A.L., Lehenbauer, T.W., Aly, S.S., 2019. Regional management 51st Annual Conference of the American Association of Bovine Practitioners, 13-
practices and prevalence of bovine respiratory disease in California’s 15 September 2018, Phoenix, AZ, p. 283.
preweaned dairy calves. Journal of Dairy Science 102, 7583–7596. NAHMS 1996. Part I: Reference of 1996 Dairy Management Practices. USDA:APHIS:
Khan, M.A., Weary, D.M., Von, K.M., 2011. Invited review: effects of milk ration on VS, Fort Collins, CO, USA.
solid feed intake, weaning, and performance in dairy heifers. Journal of Dairy NAHMS National Animal Health Monitoring System 2007. Dairy 2007 Heifer Calf
Science 94, 1071–1081. Health and Management Practices on U.S. Dairy Operations, 2007. USDA:APHIS:
King, A., Chigerwe, M., Barry, J., Murphy, J.P., Rayburn, M.C., Kennedy, E., 2020. Short VS:CEAH, Fort Collins, CO, USA.
communication: effect of feeding pooled and nonpooled high-quality colostrum Nath, M., Woolliams, J.A., Bishop, S.C., 2008. Assessment of the dynamics of
on passive transfer of immunity, morbidity, and mortality in dairy calves. microparasite infections in genetically homogeneous and heterogeneous
Journal of Dairy Science 103, 1894–1899. populations using a stochastic epidemic model. Journal of Animal Science 86,
Knauer, W.A., Godden, S.M., Dietrich, A., James, R.E., 2017. The association between 1747–1757.
daily average feeding behaviors and morbidity in automatically fed group- Neibergs, H.L., Seabury, C.M., Wojtowicz, A.J., Wang, Z., Scraggs, E., Kiser, J.N.,
housed preweaned dairy calves. Journal of Dairy Science 100, 5642–5652. Neupane, M., Womack, J.E., Van, E.A., Hagevoort, G.R., Lehenbauer, T.W., Aly, S.,
Koeck, A., Miglior, F., Kelton, D.F., Schenkel, F.S., 2012. Health recording in Canadian Davis, J., Taylor, J.F., Bovine, T., Disease, R., Coordinated, C., 2014. Susceptibility
Holsteins: data and genetic parameters. Journal of Dairy Science 95, 4099–4108. loci revealed for bovine respiratory disease complex in pre-weaned Holstein
Leblanc, S.J., Lissemore, K.D., Kelton, D.F., Duffield, T.F., Leslie, K.E., 2006. Major calves. BMC Genomics 15, 1164.
advances in disease prevention in dairy cattle. Journal of Dairy Science 89, Nor, N.M., Steeneveld, W., Van, W.T., Mourits, M.C.M., Hogeveen, H., 2013. First-
1267–1279. calving age and first-lactation milk production on Dutch dairy farms. Journal of
Lesmeister, K.E., Heinrichs, A.J., 2004. Effects of corn processing on growth Dairy Science 96, 981–992.
characteristics, rumen development, and rumen parameters in neonatal dairy Overton, M.W., 2020. Economics of respiratory disease in dairy replacement heifers.
calves. Journal of Dairy Science 87, 3439–3450. Animal Health Research Reviews 21, 143–148.
Lewis, S.J., Heaton, K.W., 1997. Stool form scale as a useful guide to intestinal transit Peel, D.S., 2020. The effect of market forces on bovine respiratory disease.
time stool form scale as a useful guide to intestinal transit time. Scandinavian Veterinary Clinics of North America - Food Animal Practice 36, 497–508.
Journal of Gastroenterology 32, 920–924. Pempek, J., Trearchis, D., Masterson, M., Habing, G., Proudfoot, K., 2017. Veal calf
Lipschutz-Powell, D., Woolliams, J.A., 2012. Bias, accuracy, and impact of indirect health on the day of arrival at growers in Ohio. Journal of Animal Science 95,
genetic effects in infectious diseases. Frontiers in Genetics 3, 215. 3863–3872.
Lipschutz-Powell, D., Woolliams, J.A., Bijma, P., Doeschl-wilson, A.B., 2012. Indirect Placzek, M., Christoph-Schulz, I., Barth, K., 2021. Public attitude towards cow-calf
genetic effects and the spread of infectious disease: are we capturing the full separation and other common practices of calf rearing in dairy farming—a
heritable variation underlying disease prevalence? PLoS One 7, e39551. review. Organic Agriculture 11, 41–50.
Lipschutz-Powell, D., Woolliams, J.A., Doeschl-Wilson, A.B., 2013. A unifying theory Pritchett, L.C., Gay, C.C., Besser, T.E., Hancock, D.D., 1991. Management and
for genetic epidemiological analysis of binary disease data. Genetics Selection production factors influencing immunoglobulin G1 concentration in
Evolution 46, 1–12. colostrum from Holstein cows. Journal of Dairy Science 74, 2336–2341.
Lipschutz-Powell, D., Woolliams, J.A., Doeschl-Wilson, A.B., 2014. A unifying theory Pro Earth Animal Health 2017. Part III: Calf Scours The Economic Impact. Pro Earth
for genetic epidemiological analysis of binary disease data. Genetics Selection Animal Health. Retrieved on 9 January 2023 from https://
Evolution 46, 1–12. proearthanimalhealth.com/wp-content/uploads/2018/02/Calf-Scours-
Lombard, J.E., Garry, F.B., Tomlinson, S.M., Garber, L.P., 2007. Impacts of Economic-ImpactsPDF.pdf.
dystocia on health and survival of dairy calves. Journal of Dairy Science 90, Probo, M., Veronesi, M.C., 2022. Clinical scoring systems in the newborn calf: an
1751–1760. overview. Animals 12, 3013.
Love, W.J., Lehenbauer, T.W., Kass, P.H., Van Eenennaam, A.L., Aly, S.S., 2014. Quick, A.E., Ollivett, T.L., Kirkpatrick, B.W., Weigel, K.A., 2020. Genomic analysis of
Development of a novel clinical scoring systemfor on-farmdiagnosis of bovine bovine respiratory disease and lung consolidation in preweaned Holstein calves
respiratory disease in pre-weaned dairy calves. PeerJ 2, e238. using clinical scoring and lung ultrasound. Journal of Dairy Science 103, 1632–
Lynch, C., Schenkel, F.S., van Staaveren, N., Miglior, F., Kelton, D., Baes, C.F., 2024. 1641.
Investigating the potential for genetic selection of dairy calf disease traits using Quigley, J.D., Lago, A., Chapman, C., Erickson, P., Polo, J., 2013. Evaluation of the Brix
management data. Journal of Dairy Science 107, 1022–1034. refractometer to estimate immunoglobulin G concentration in bovine
Mahmoud, M., Yin, T., Brügemann, K., König, S., 2017. Phenotypic, genetic, and colostrum. Journal of Dairy Science 96, 1148–1155.
single nucleotide polymorphism marker associations between calf diseases and Raboisson, D., Trillat, P., Cahuzac, C., 2016. Failure of passive immune transfer in
subsequent performance and disease occurrences of first-lactation German calves: a meta-analysis on the consequences and assessment of the economic
Holstein cows. Journal of Dairy Science 100, 2017–2031. impact. PLoS ONE 11, 1–19.
Malchiodi, F., Koeck, A., Mason, S., Christen, A.M., Kelton, D.F., Schenkel, F.S., Miglior, Renaud, D.L., Kelton, D.F., LeBlanc, S.J., Haley, D.B., Duffield, T.F., 2018. Calf
F., 2017. Genetic parameters for hoof health traits estimated with linear and management risk factors on dairy farms associated with male calf mortality
threshold models using alternative cohorts. Journal of Dairy Science 100, 2828– on veal farms. Journal of Dairy Science 101, 1785–1794.
2836. Renaud, D.L., Buss, L., Wilms, J.N., Steele, M.A., 2020. Technical note: is fecal
Martin, P., Vinet, A., Denis, C., Grohs, C., Chanteloup, L., Dozias, D., Maupetit, D., Sapa, consistency scoring an accurate measure of fecal dry matter in dairy calves?
J., Renand, G., Blanc, F., 2021. Determination of immunoglobulin concentrations Journal of Dairy Science 103, 10709–10714.

10
C. Lynch, E.M. Leishman, F. Miglior et al. Animal 18 (2024) 101141

Rivero, M.J., Valderrama, X., Haines, D., Alomar, D., 2012. Prediction of USDA 2018. Dairy 2014: Health and Management Practices on U.S. Dairy
immunoglobulin G content in bovine colostrum by near-infrared Operations, 2014. Retrieved on 22 November 2022, from https://www.aphis.
spectroscopy. Journal of Dairy Science 95, 1410–1418. usda.gov/animal_health/nahms/dairy/downloads/dairy14/Dairy14_dr_PartIII.
Robbers, L., Jorritsma, R., Nielen, M., Koets, A., 2021. A scoping review of on-farm pdf.
colostrum management practices for optimal transfer of immunity in dairy Uyama, T., Renaud, D.L., Morrison, E.I., McClure, J.T., LeBlanc, S.J., Winder, C.B., de
calves. Frontiers in Veterinary Science 8, 1–16. Jong, E., McCubbin, K.D., Barkema, H.W., Dufour, S., Sanchez, J., Heider, L.C.,
Roche, S.M., Von Massow, M., Renaud, D., Shock, D.A., Jones-Bitton, A., Kelton, D.F., Kelton, D.F., 2022. Associations of calf management practices with
2020. Cost-benefit of implementing a participatory extension model for antimicrobial use in Canadian dairy calves. Journal of Dairy Science 105,
improving on-farm adoption of Johne’s disease control recommendations. 9084–9097.
Journal of Dairy Science 103, 451–472. van Staaveren, N., Hyland, E., Houlahan, K., Lynch, C., Miglior, F., Kelton, D.F.,
Roff, D.A., 1996. The evolution of threshold traits in animals. The Quarterly Review Schenkel, F.S., Baes, C.F., 2023. Recording of calf diseases for potential use in
of Biology 71, 3–35. breeding programs: a case study on calf respiratory illness and diarrhea.
Roso, V.M., Schenkel, F.S., Miller, S.P., 2004. Degree of connectedness among Canadian Journal of Animal Science 103, 192–203.
groups of centrally tested beef bulls. Canadian Journal of Animal Science 84, 37– VanRaden, P.M., Van Tassell, C.P., Wiggans, G.R., Sonstegard, T.S., Schnabel, R.D.,
47. Taylor, J.F., Schenkel, F.S., 2009. Invited review: reliability of genomic
RSPCA 2021. RSPCA welfare standards for Dairy cattle. RSPCA. Retrieved on 21 predictions for North American Holstein bulls. Journal of Dairy Science 92,
November 2022, from https://science.rspca.org.uk/documents/1494935/ 16–24.
9042554/RSPCA+Dairy+Standards.pdf/bb1a2971-8018-b325-8dcc- Ventura, B.A., Von, K.M., Schuppli, C.A., Weary, D.M., 2013. Views on contentious
d6b590fcd955?t=1625822966824. practices in dairy farming : the case of early cow-calf separation. Journal of
Santman-Berends, I.M.G.A., Schukken, Y.H., van Schaik, G., 2019. Quantifying calf Dairy Science 96, 6105–6116.
mortality on dairy farms: challenges and solutions. Journal of Dairy Science 102, Virtala, A.K., Mechor, G.D., Grohn, Y.T., 1996. The effect of calfhood diseases on
6404–6417. growth of female dairy calves during the first 3 months of life in New York state.
Schefers, J.M., Weigel, K.A., 2012. Genomic selection in dairy cattle: integration of Journal of Dairy Science 79, 1040–1049.
DNA testing into breeding programs. Animal Frontiers 2, 4–9. Waltner-Toews, D., Martin, S., Meek, A., 1986. The effect of early calfhood health
Schinwald, M., Creutzinger, K., Winder, C.B., Haley, D., Renaud, D.L., 2022. Predictors status on survivorship and age at first calving. Canadian Journal of Veterinary
of diarrhea, mortality, and weight gain in male dairy calves. Journal of Dairy Research 50, 314–317.
Science 105, 5296–5309. Wathes, D.C., Brickell, J.S., Bourne, N.E., Swali, A., Cheng, Z., 2008. Factors influencing
Shook, G.E., 1989. Selection for disease resistance. Journal of Dairy Science 72, heifer survival and fertility on commercial dairy farms. Animal 2, 1135–1143.
1349–1362. Weaver, D.M., Tyler, J.W., Vanmetre, D.C., Hostetler, D.E., Barrington, G.M., 2000.
Silva FG, Lamy E, Pedro S, Azevedo I, Caetano P, Ramalho J, Martins L, Pereira AMF, Passive transfer of colostral immunoglobulins in calves. Journal of Veterinary
Cerqueira JOL, Silva SR and Conceição C 2023. Assessing passive immune Internal Medicine 14, 569–577.
transfer in newborn calves: salivary and serum IgG association. Proceedings of Wiggans, G.R., Carrillo, J.A., 2022. Genomic selection in United States dairy cattle.
the 74th EAAP Annual Meeting, 26 August – 1 September 2023, Lyon, France, p. Frontiers in Genetics 13, 1–7.
77. Wiggans, G.R., Cole, J.B., Hubbard, S.M., Sonstegard, T.S., 2017. Genomic selection in
Sischo, W.M., Hird, D.W., Gardner, I.A., Utterback, W.W., Christiansen, K.H., dairy cattle: the USDA experience. Annual Review of Animal Biosciences 5, 309–
Carpenter, T.E., Danaye-Elmi, C., Heron, B.R., 1990. Economics of disease 329.
occurrence and prevention on California dairy farms: a report and evaluation Williams, D.R., Pithua, P., Garcia, A., Champagne, J., Haines, D.M., Aly, S.S., 2014.
of data collected for the national animal health monitoring system, 1986–87. Effect of three colostrum diets on passive transfer of immunity and preweaning
Preventive Veterinary Medicine 8, 141–156. health in calves on a California dairy following colostrum management training.
Sivula, N.J., Ames, T.R., Marsh, W.E., Werdin, R.E., 1996. Descriptive epidemiology of Veterinary Medicine International 2014, 1–9.
morbidity and mortality in Minnesota dairy heifer calves. Preventive Veterinary Wilson, D.J., Habing, G., Winder, C.B., Renaud, D.L., 2023. A scoping review of
Medicine 27, 155–171. neonatal calf diarrhea case definitions. Preventive Veterinary Medicine 211,
Sorge, U., Kelton, D., Staufenbiel, R., 2009. Neonatal blood lactate concentration and 105818.
calf morbidity. Veterinary Record 164, 553–554. Winder, C.B., Bauman, C.A., Duffield, T.F., Barkema, H.W., Keefe, G.P., Dubuc, J.,
Soufleri, A., Banos, G., Panousis, N., Fletouris, D., Arsenos, G., Valergakis, G.E., 2019. Uehlinger, F., Kelton, D.F., 2018. Canadian national dairy study: heifer calf
Genetic parameters of colostrum traits in Holstein dairy cows. Journal of Dairy management. Journal of Dairy Science 101, 10565–10579.
Science 102, 11225–11232. Windeyer, M.C., Leslie, K.E., Godden, S.M., Hodgins, D.C., Lissemore, K.D., LeBlanc, S.
Springer, H., Yost, H., 2022. Penn state extension: neonatal calf diarrhea (Scours). J., 2014. Factors associated with morbidity, mortality, and growth of dairy heifer
The Pennsylvania State University, State College, PA, USA. calves up to 3 months of age. Preventive Veterinary Medicine 113, 231–240.
Stanton A 2011. An evaluation of the impact of management practice on the health Wolfger, B., Jones, B.W., Orsel, K., Bewley, J.M., 2017. Technical note : evaluation of
and welfare of dairy Heifer calves. University of Guelph. Retrieved on 21 an ear-attached real-time location monitoring system. Journal of Dairy Science
November 2022, from https://atrium.lib.uoguelph.ca/xmlui/handle/10214/ 100, 2219–2224.
2856. Woolums, A., 2021. Vaccination protocols for dairy calves. American Association of
Stilwell, G., Carvalho, R.C., 2011. Clinical outcome of calves with failure of passive Bovine Practioners 54, 178–180.
transfer as diagnosed by a commercially available IgG quick test kit. Canadian Yin, T., Halli, K., König, S., 2022. Direct genetic effects, maternal genetic effects, and
Veterinary Journal 52, 524–526. maternal genetic sensitivity on prenatal heat stress for calf diseases and
Svensson, C., Hultgren, J., 2008. Associations between housing, management, and corresponding genomic loci in German Holsteins. Journal of Dairy Science 105,
morbidity during rearing and subsequent first-lactation milk production of 6795–6808.
dairy cows in Southwest Sweden. Journal of Dairy Science 91, 1510–1518. Zhang, W., Chang, L., Brito, D., Shi, W., Dong, L., 2019b. Mortality-Culling rates of
Taylor, J.D., Fulton, R.W., Lehenbauer, T.W., Step, D.L., Confer, A.W., 2010. The dairy calves and replacement heifers and its risk factors in Holstein Cattle.
epidemiology of bovine respiratory disease: what is the evidence for Animals 9, 730.
predisposing factors? Canadian Veterinary Journal 51, 1095–1102. Zhang, Z., Kargo, M., Liu, A., Thomasen, J.R., Pan, Y., Su, G., 2019a. Genotype-by-
Tora, E., Abayneh, E., Seyoum, W., Shurbe, M., 2021. Longitudinal study of calf environment interaction of fertility traits in Danish Holstein cattle using a
morbidity and mortality on smallholder farms in southern Ethiopia. PLoS ONE single-step genomic reaction norm model. Heredity 123, 202–214.
16, 1–18. Zhang, H., Wang, K., An, T., Zhu, L., Chang, Y., Lou, W., Liu, A., Su, G., Brito, L.F., Wang,
Tsairidou, S., Anacleto, O., Woolliams, J.A., Doeschl-Wilson, A., 2019. Enhancing Y., 2022. Genetic parameters for dairy calf and replacement heifer wellness
genetic disease control by selecting for lower host infectivity and susceptibility. traits and their association with cow longevity and health indicators in Holstein
Heredity 122, 742–758. cattle. Journal of Dairy Science 105, 6749–6759.
Urie, N.J., Lombard, J.E., Shivley, C.B., Kopral, C.A., Adams, A.E., Earleywine, T.J., Zoetis 2022. Dairy Wellness Profit IndexÒ enhancements for profitable genetic
Olson, J.D., Garry, F.B., 2018. Preweaned heifer management on US dairy selection and helping sustainable practices. Zoetis. Retrieved on 9 January 2023
operations: part I. descriptive characteristics of preweaned heifer raising from https://www.zoetisus.com/content/pages/Solutions/Dairy/Dairy-
practices. Journal of Dairy Science 101, 9168–9184. Genetics/Dairy-genetics-resources/Documents/Genetics-edetailer.pdf.

11