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A cattle family in New Zealand with triplet calving ability
Livestock Science 128 (2010) 193–196
Contents lists available at ScienceDirect
Livestock Science j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / l i v s c i
Short communication
A cattle family in New Zealand with triplet calving ability C.A. Morris a,⁎, M. Wheeler a, G.L. Levet b, B.W. Kirkpatrick c a b c
AgResearch, Ruakura Research Centre, PB 3123, Hamilton 3240, New Zealand Kikitangeo, Hoteo North, RD2 Wellsford, New Zealand Department of Animal Sciences, University of Wisconsin, Madison, Wisconsin 53706, USA
a r t i c l e
i n f o
Article history: Received 23 August 2009 Received in revised form 25 November 2009 Accepted 26 November 2009 Keywords: Cow Triplet Inheritance
a b s t r a c t This paper reports on calving data from a cow family in New Zealand which is elite for triplet calving ability. The foundation cow (named ‘Treble’) was born in 1993 and her calvings included three sets of triplets. A triplet son (named ‘Trio’), born in 1996, was used for matings in two New Zealand herds, producing 44 daughters which were retained for breeding, and these had a total of 109 calvings by 2008. Thirteen of the daughters produced a total of 15 twin and 6 triplet sets, 19% multiples (including twins and triplets). Five cows calved the 6 triplet sets; triplet calvings were 29% of all multiple calvings. This herd should provide a resource for DNA studies to identify a genetic mechanism whereby rare triplet calvings are produced. © 2009 Elsevier B.V. All rights reserved.
1. Introduction Twin and triplet births are uncommon in cattle. In New Zealand dairy cattle (mainly Friesians, Jerseys and their crosses), 1.0% of calvings result in twins, with a range by geographical region from 0.8 to 1.4% (New Zealand Dairy Board, 1961). In New Zealand pedigree beef cattle, 0.98% of calvings overall result in twins, with upper values of 1.7% in the Charolais and 2.1% in the Simmental breeds (Morris and Packard, 1985). In the USA, Rutledge (1975) reported average twin calving rates by breed, including Holstein (3.4%), Jersey (1.3%), Angus (1.1%) and Hereford (0.4%). A high twinning incidence is often considered undesirable in cattle (e.g., Nielen et al., 1989), although generally desirable in sheep (Gordon, 2004), and understanding the underlying biology of non-identical twin and triplet production in cattle may assist in any attempts to breed for or against the trait. Generally, twinning rate in cattle has been found to have a low heritability, 0.10 or less (e.g. Syrstad, 1984; Gregory et al., 1997; Karlsen et al., 2000) but, in spite of this, long-term experimental selection for increased twinning rate has been
⁎ Corresponding author. Tel.: +64 78385417; fax: +64 78385038. E-mail address: [email protected] (C.A. Morris). 1871-1413/$ – see front matter © 2009 Elsevier B.V. All rights reserved. doi:10.1016/j.livsci.2009.11.009
successful (e.g., Morris and Wheeler, 2002; Echternkamp et al., 2007a). Both of the twin-selection herds cited have involved screening industry herds intensely for outlier animals in order to establish the foundation populations in New Zealand and at the US Department of Agriculture, Clay Center, Nebraska, respectively. Such herds are known to comprise cows calving non-identical twin sets (e.g., as deduced from the sex distributions of 772 twin-pairs (Morris, 1991), and from twin ovulation studies (Echternkamp et al., 2007a)). Relative to twin calvings, triplet calvings are even less common, with the rarity depending on mean twin incidence, as described originally for human multiple births by Hellin's Law (1895). In New Zealand dairy industry data (H. J. Tempero, Livestock Improvement Corporation, Hamilton, New Zealand, personal communication: 1995), triplet calvings in Holstein–Friesians from 1991/92 to 1994/95 occurred at 0.0071 of the rate of twin calvings. In contrast, in a high twinning rate environment in the American experimental population cited above where selection was for increased numbers of multiples (defined as twins or higher order births), Bennett et al. (2006) reported that the incidence of triplet calvings was 0.081 of the incidence of twin calvings (a figure more than 11 times higher than in commercial New Zealand dairy cattle).
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C.A. Morris et al. / Livestock Science 128 (2010) 193–196
The present report describes an elite cow family in New Zealand with a high incidence of triplet calvings, which should provide a breeding or DNA resource for further study.
and subsequently these had calvings of their own (described below). 2.5. Data analyses
2. Materials and methods 2.1. Ethics This research was carried out with the approval of the AgResearch Ruakura Animal Ethics Committee.
Annual calving data (with single calvings, or multiples (twins or triplets)) were analysed, and summarised according to source (AgResearch selected herd, or Levet unselected herd). 3. Results
2.2. Foundation cow 3.1. Daughters born at AgResearch in 2000 and 2001 A highly fecund cow was identified in 1996 by one of us (GLL) in his beef herd, located near Wellsford, New Zealand. Subsequently the cow was given the name ‘Treble’, and up to 1996 she had calved two sets of triplets. Treble's history and calving record was: • Born in 1993, and purchased as a calf at foot alongside her dam. No other detailed history is available. • She is of uncertain breed, but her coat colours are indicative of at least a three-breed cross, possibly Hereford × (Friesian× Jersey). • Her dam calved a set of twins in 2000. • Treble calved her first triplet set as a two-year-old in 1995, a live heifer and two dead calves of unrecorded sex. • She calved her second set in late 1996, three live calves (two heifers and a bull). The bull was named ‘Trio’, and has been used for natural matings, described below, and for semen collection. He was culled in 2008 at 12 years of age. • Treble carried a third triplet set to term in April 1999, but the calves were delivered with considerable difficulty and veterinary assistance. All calves were dead, and sexes were not recorded. • Treble then became infertile, apparently as a result of the calving difficulty in 1999; she was culled in 2007 at 14 years of age. • She has been cloned, and two clone ‘offspring’ were born at AgResearch's Ruakura Research Centre, Hamilton, in November 2000. 2.3. Matings by Trio at AgResearch Trio was mated to selected cows on an AgResearch farm to produce calves in 2000 and 2001. The cows (mainly Friesian and Friesian-crosses, with some Charolais- and Simmentalcrosses) were part of the twinning rate selection herd developed since 1983/84 by AgResearch, a herd with an average annual twin calving rate which had risen to 13.3% by 1998–2000 (Morris and Wheeler, 2002); this included no triplet calvings. There were 14 resulting daughters of Trio, and subsequently these had calvings of their own (described below). 2.4. Matings by Trio on the Levet property Trio was mated in the Levet herd to Friesian-cross cows which were unrecorded (but not noticeably different in twinning rate from ∼1 to 2% for the district), to produce calves in 2002–04. There were 30 resulting daughters of Trio,
Fourteen daughters of Trio calved on an AgResearch farm until 2003, after which time most were transferred to the Levet property. Their calving performance is summarized in Table 1, subdivided into calving results up to, or after, 2003. They had 56 calvings, including five multiples (four twin sets and one triplet set) from three cows. 3.2. Daughters born on the Levet property in 2002–2004 Table 1 shows the calving performance of 30 daughters of Trio, born on the Levet property. Two, four and four of these daughters, born in 2002, 2003 and 2004, respectively, calved multiples (a total of 11 twin sets and 5 triplet sets). 3.3. All daughters of Trio Combining all birth cohorts of Trio's daughters (Table 1), 13 cows (30%) out of his total of 44 daughters produced at least one multiple, with 21 (19%) out of 109 calvings being multiple births, and 6 sets (29%) out of 21 multiples being triplet sets. For those cows calving triplets, surprisingly two sets were produced by 2-year-olds, two sets by 3-year-olds, and the one cow with two sets calved these as a 3-year-old and a 4-year-old. 3.4. Calf survival Calf survival for the triplet-born calves from Trio's daughters was low. Eight calves (44%) survived from six sets. One of these sets was premature (all died). For the other five sets, denoting A = born alive and D = born dead, there were two AAA sets, two DDD sets and one AAD. No post mortem investigations were carried out. Adding data from the original three sets produced by Treble herself, 12 calves (also 44%) survived from nine sets. 3.5. Associated animals and data 1. Blood samples for DNA were collected in 2006 from 42 daughters of Trio, for molecular study at the University of Wisconsin, USA. DNA has also been stored from Treble herself. 2. Semen was collected from Trio in 2006, and exported to Wisconsin for one of us (BWK) to generate more daughters by artificial insemination and embryo transplant. A total of 36 Trio daughters were born in the USA in 2008 and are currently being evaluated as yearlings for ovulation rate.
C.A. Morris et al. / Livestock Science 128 (2010) 193–196
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Table 1 Calving history up to 2008 for ‘Trio’-sired cows born at AgResearch and on the Levet property. Birth location Year of birth
Calving years
Calving location Number of cows Cows with multiples Number of calvings b Twin sets Triplet sets
AgResearch
2000 and 2001 2002 and 2003 AgResearch 2004–07 AgResearch 2004–08 Levet
Levet
2002 2003 2004
Total
2004–08 2005–08 2006–08
Levet Levet Levet
14 a (1) (10) 14 4 10 16 30 44
3a (1) (0) 3 2 4 4 10 13
18 3 35 56 10 19 24 53 109
3 1 0 4 2 5 4 11 15
1 0 0 1 1 2 2 5 6
a
The 14 cows born at AgResearch and calving at AgResearch up to 2003 had post-2003 calvings in two locations, shown in the subsequent two rows (three other cows did not calve again). b Includes multiples (combining twin and triplet sets) and single calvings.
Approximately 55 and 35 additional Trio daughters are expected to be born in 2009 and 2010, respectively. All these offspring will be used for ovulation research and molecular genetic studies. 3. Nuclear transfer was performed with skin fibroblasts obtained from an ear punch from Treble, and two clones were generated in 2000 (D.N. Wells, AgResearch, personal communication: 2000). At ∼ 18 months of age, these clones were observed with triplet and quadruplet ovulations after normal oestrous patterns, although triplet and quadruplet ovulations are otherwise a rare phenomenon in cattle. By 2007, one of the clones produced only one set of twins. 4. Discussion This report describes an elite cow, Treble, which calved three sets of triplets, and had exceptional reproductive performance in her descendants, especially six sets of triplets produced by some of her 44 grand-daughters descended through one son, Trio. The high incidence of twin- and tripletproducing animals (13 (30%) of all grand-daughters) was consistent with the high performance of some prolific cow families resulting from nearly 20 years of screening and selection for twinning in an AgResearch herd (Morris and Wheeler, 2002), but with the additional attribute of triplet calvings (six (29%) out of 21 multiple births were triplet sets). In the experimental twinning herd at Clay Center, Nebraska, Echternkamp et al. (2007a) found that cows recorded with 1, 2 or 3 foetuses had, on average, 95.7 ± 0.6%, 87.8 ± 0.8% and 54.9 ± 2.3% pregnancies maintained to term, respectively. Echternkamp et al. (2007b) suggested that triplet pregnancies in cattle were compromised in dams of all ages, by physical or physiological limitations. It is therefore suggested that cows in Treble's family may have not only high ovulation rates but also sufficient uterine capacity to carry calves to term. If this is the result of a mutation of a major gene for ovulation rate, this mutation would need to be present in a background which provides for high uterine capacity, and that background would still need to be sufficient when diluted to 25% in grand-daughters. For single-gene inheritance, a gene or gene complex would need to be transmitted as a single copy from Treble to some granddaughters via a son, and such a gene would also need to be expressed in females as a dominant or partially dominant.
The data on Trio's daughters described in Table 1 (30% producing at least one multiple, 19% of all calvings being multiple births, and 29% of the multiples being triplet sets) can be compared with: A. the experimental twinning herd at AgResearch which had relatively high twinning rates (excluding the Trio family, there were 33% twin births in six calving years, 2003–08; C.A. Morris, unpublished data: 2009) but no triplets, and B. other cows on the Levet property derived from the AgResearch high twinning rate herd, without any known ancestral link to Trio or Treble, and their figures were 14 sets of twins (12.6%) from 111 calvings, and no triplets. The cow performance details in Table 1 needed to be separated into cow-birth years because the cows had different groups of foundation dams, from the AgResearch twin-breeding herd or the unselected Levet herd. Attributing the triplet incidence to a cow-sire effect might be uncertain in the case of those derived from AgResearch dams because of their experimental selection history; however, in 26 years in the AgResearch herd, there have been no other cases of triplets, from 953 cows with 2677 calvings. Some other exceptional cows or cow families have been reported in 10 historical references reviewed by Morris (1984), including a ‘Black Polled’ cow with 25 calves from eight calvings (one single, two sets each of twins, triplets and quadruplets, and a set of sextuplets), and a cow with 15 calves from five calvings (a single and two sets each of triplets and quadruplets). In the Clay Center, Nebraska, herd above and in North American Holsteins, Kim et al. (2009) have reported successful genomic searches for quantitative trait loci (QTL) on bovine chromosome (BTA) 5 for multiple births, and a Norwegian study has shown QTL on BTA 5, 7, 12 and 23 (Lien et al., 2000). Other QTL locations have also been reported by Arias and Kirkpatrick (2004) and by Cobanoglu et al. (2005). The Treble family should provide the basis for a similar genomic study, including DNA samples from Trio's daughters in the Levet herd, New Zealand, and also from Trio's daughters born in the USA following the importation of his semen there. In conclusion, Treble is a cow of remarkable reproductive performance, with three sets of triplets. The fact that some of her grand-daughters also produced triplets strongly suggests that some of her ability is inherited. Treble and her family should provide a valuable resource for further study of variation in bovine fecundity.
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Acknowledgements We thank the AgResearch Farm Managers and staff, and Mr. Alan Piggott (Levet herd), for care of stock and for collection of calving data. References Arias, J.A., Kirkpatrick, B.W., 2004. Mapping of bovine ovulation rate QTL; an analytical approach for three generation pedigrees. Anim. Genet. 35, 7–13. Bennett, G.L., Allan, M.F., Cushman, R.A., Echternkamp, S.E., 2006. Observed and predicted numbers of single, twin, and triplet births in a cattle population selected for increased twinning. J. Anim. Sci. 84 (Suppl. 1) Abstr. T135. Cobanoglu, O., Berger, P.J., Kirkpatrick, B.W., 2005. Genome screen for twinning rate QTL in four North American Holstein families. Anim. Genet. 36, 303–308. Echternkamp, S.E., Cushman, R.A., Allan, M.F., Thallman, R.M., Gregory, K.E., 2007a. Effects of ovulation rate and fetal number on fertility in twinproducing cattle. J. Anim. Sci. 85, 3228–3238. Echternkamp, S.E., Thallman, R.M., Cushman, R.A., Allan, M.F., Gregory, K.E., 2007b. Increased calf production in cattle selected for twin ovulations. J. Anim. Sci. 85, 3239–3248. Gordon, I.R., 2004. Controlling multiple births and litter size. Reproductive Technologies in Farm Animals. CABI Publishing, pp. 195–213. Chapter 8. Gregory, K.E., Bennett, G.L., Van Vleck, L.D., Echternkamp, S.E., Cundiff, L.V., 1997. Genetic and environmental parameters for ovulation rate, twinning rate, and weight traits in a cattle population selected for twinning. J. Anim. Sci. 75, 1213–1222.
Hellin, D., 1895. Die Ursache der Multiparität der uniparen Tiere überhaupt und der Zwillings-schwangerschaft beim Menschen insbesondere. Seitz & Schauer, München, pp. 1–70. Karlsen, A., Ruane, J., Klemetsdal, G., Heringstad, B., 2000. Twinning rate in Norwegian cattle: frequency, (co)variance components, and genetic trends. J. Anim. Sci. 78, 15–20. Kim, E.-S., Shi, X., Cobanoglu, O., Weigel, K., Berger, P.J., Kirkpatrick, B.W., 2009. Refined mapping of twinning-rate quantitative trait loci on bovine chromosome 5 and analysis of insulin-like growth factor-1 as a positional candidate gene. J. Anim. Sci. 87, 835–843. Lien, S., Karlsen, A., Klemetsdal, G., Vage, D.I., Olsaker, I., Klungland, H., Aasland, M., Heringstad, B., Ruane, J., Gomez-Raya, L., 2000. A primary screen of the bovine genome for quantitative trait loci affecting twinning rate. Mamm. Genome 11, 877–882. Morris, C.A., 1984. A review of the genetics and reproductive physiology of dizygotic twinning in cattle. Anim. Breed. Abstr. 52, 803–819. Morris, C.A., 1991. Screening herds for cows with a history of twin calving. Proc. N. Z. Soc. Anim. Prod. 51, 447–451. Morris, C.A., Packard, P.M., 1985. Progress with Beefplan. Annual Report 1984–85, Ruakura Animal Research Station (Genetics Section). Ministry of Agriculture & Fisheries, Hamilton, New Zealand, pp. 76–77. Morris, C.A., Wheeler, M., 2002. Genetic variation in twin calving incidence in herds with a high phenotypic mean. N. Z. J. Agric. Res. 45, 17–25. New Zealand Dairy Board, 1961. 37th Farm Production Report (Table 4). Farm Production Division, Wellington, New Zealand, p. 78. Nielen, M., Schukken, Y.H., Scholl, D.T., Wilbrink, H.J., Brand, A., 1989. Twinning in dairy cattle: a study of risk factors and effects. Theriogenology 32, 845–862. Rutledge, J.J., 1975. Twinning in cattle. J. Anim. Sci. 40, 803–815. Syrstad, O., 1984. Inheritance of multiple births in cattle. Livest. Prod. Sci. 11, 373–380.
Contents lists available at ScienceDirect
Livestock Science j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / l i v s c i
Short communication
A cattle family in New Zealand with triplet calving ability C.A. Morris a,⁎, M. Wheeler a, G.L. Levet b, B.W. Kirkpatrick c a b c
AgResearch, Ruakura Research Centre, PB 3123, Hamilton 3240, New Zealand Kikitangeo, Hoteo North, RD2 Wellsford, New Zealand Department of Animal Sciences, University of Wisconsin, Madison, Wisconsin 53706, USA
a r t i c l e
i n f o
Article history: Received 23 August 2009 Received in revised form 25 November 2009 Accepted 26 November 2009 Keywords: Cow Triplet Inheritance
a b s t r a c t This paper reports on calving data from a cow family in New Zealand which is elite for triplet calving ability. The foundation cow (named ‘Treble’) was born in 1993 and her calvings included three sets of triplets. A triplet son (named ‘Trio’), born in 1996, was used for matings in two New Zealand herds, producing 44 daughters which were retained for breeding, and these had a total of 109 calvings by 2008. Thirteen of the daughters produced a total of 15 twin and 6 triplet sets, 19% multiples (including twins and triplets). Five cows calved the 6 triplet sets; triplet calvings were 29% of all multiple calvings. This herd should provide a resource for DNA studies to identify a genetic mechanism whereby rare triplet calvings are produced. © 2009 Elsevier B.V. All rights reserved.
1. Introduction Twin and triplet births are uncommon in cattle. In New Zealand dairy cattle (mainly Friesians, Jerseys and their crosses), 1.0% of calvings result in twins, with a range by geographical region from 0.8 to 1.4% (New Zealand Dairy Board, 1961). In New Zealand pedigree beef cattle, 0.98% of calvings overall result in twins, with upper values of 1.7% in the Charolais and 2.1% in the Simmental breeds (Morris and Packard, 1985). In the USA, Rutledge (1975) reported average twin calving rates by breed, including Holstein (3.4%), Jersey (1.3%), Angus (1.1%) and Hereford (0.4%). A high twinning incidence is often considered undesirable in cattle (e.g., Nielen et al., 1989), although generally desirable in sheep (Gordon, 2004), and understanding the underlying biology of non-identical twin and triplet production in cattle may assist in any attempts to breed for or against the trait. Generally, twinning rate in cattle has been found to have a low heritability, 0.10 or less (e.g. Syrstad, 1984; Gregory et al., 1997; Karlsen et al., 2000) but, in spite of this, long-term experimental selection for increased twinning rate has been
⁎ Corresponding author. Tel.: +64 78385417; fax: +64 78385038. E-mail address: [email protected] (C.A. Morris). 1871-1413/$ – see front matter © 2009 Elsevier B.V. All rights reserved. doi:10.1016/j.livsci.2009.11.009
successful (e.g., Morris and Wheeler, 2002; Echternkamp et al., 2007a). Both of the twin-selection herds cited have involved screening industry herds intensely for outlier animals in order to establish the foundation populations in New Zealand and at the US Department of Agriculture, Clay Center, Nebraska, respectively. Such herds are known to comprise cows calving non-identical twin sets (e.g., as deduced from the sex distributions of 772 twin-pairs (Morris, 1991), and from twin ovulation studies (Echternkamp et al., 2007a)). Relative to twin calvings, triplet calvings are even less common, with the rarity depending on mean twin incidence, as described originally for human multiple births by Hellin's Law (1895). In New Zealand dairy industry data (H. J. Tempero, Livestock Improvement Corporation, Hamilton, New Zealand, personal communication: 1995), triplet calvings in Holstein–Friesians from 1991/92 to 1994/95 occurred at 0.0071 of the rate of twin calvings. In contrast, in a high twinning rate environment in the American experimental population cited above where selection was for increased numbers of multiples (defined as twins or higher order births), Bennett et al. (2006) reported that the incidence of triplet calvings was 0.081 of the incidence of twin calvings (a figure more than 11 times higher than in commercial New Zealand dairy cattle).
194
C.A. Morris et al. / Livestock Science 128 (2010) 193–196
The present report describes an elite cow family in New Zealand with a high incidence of triplet calvings, which should provide a breeding or DNA resource for further study.
and subsequently these had calvings of their own (described below). 2.5. Data analyses
2. Materials and methods 2.1. Ethics This research was carried out with the approval of the AgResearch Ruakura Animal Ethics Committee.
Annual calving data (with single calvings, or multiples (twins or triplets)) were analysed, and summarised according to source (AgResearch selected herd, or Levet unselected herd). 3. Results
2.2. Foundation cow 3.1. Daughters born at AgResearch in 2000 and 2001 A highly fecund cow was identified in 1996 by one of us (GLL) in his beef herd, located near Wellsford, New Zealand. Subsequently the cow was given the name ‘Treble’, and up to 1996 she had calved two sets of triplets. Treble's history and calving record was: • Born in 1993, and purchased as a calf at foot alongside her dam. No other detailed history is available. • She is of uncertain breed, but her coat colours are indicative of at least a three-breed cross, possibly Hereford × (Friesian× Jersey). • Her dam calved a set of twins in 2000. • Treble calved her first triplet set as a two-year-old in 1995, a live heifer and two dead calves of unrecorded sex. • She calved her second set in late 1996, three live calves (two heifers and a bull). The bull was named ‘Trio’, and has been used for natural matings, described below, and for semen collection. He was culled in 2008 at 12 years of age. • Treble carried a third triplet set to term in April 1999, but the calves were delivered with considerable difficulty and veterinary assistance. All calves were dead, and sexes were not recorded. • Treble then became infertile, apparently as a result of the calving difficulty in 1999; she was culled in 2007 at 14 years of age. • She has been cloned, and two clone ‘offspring’ were born at AgResearch's Ruakura Research Centre, Hamilton, in November 2000. 2.3. Matings by Trio at AgResearch Trio was mated to selected cows on an AgResearch farm to produce calves in 2000 and 2001. The cows (mainly Friesian and Friesian-crosses, with some Charolais- and Simmentalcrosses) were part of the twinning rate selection herd developed since 1983/84 by AgResearch, a herd with an average annual twin calving rate which had risen to 13.3% by 1998–2000 (Morris and Wheeler, 2002); this included no triplet calvings. There were 14 resulting daughters of Trio, and subsequently these had calvings of their own (described below). 2.4. Matings by Trio on the Levet property Trio was mated in the Levet herd to Friesian-cross cows which were unrecorded (but not noticeably different in twinning rate from ∼1 to 2% for the district), to produce calves in 2002–04. There were 30 resulting daughters of Trio,
Fourteen daughters of Trio calved on an AgResearch farm until 2003, after which time most were transferred to the Levet property. Their calving performance is summarized in Table 1, subdivided into calving results up to, or after, 2003. They had 56 calvings, including five multiples (four twin sets and one triplet set) from three cows. 3.2. Daughters born on the Levet property in 2002–2004 Table 1 shows the calving performance of 30 daughters of Trio, born on the Levet property. Two, four and four of these daughters, born in 2002, 2003 and 2004, respectively, calved multiples (a total of 11 twin sets and 5 triplet sets). 3.3. All daughters of Trio Combining all birth cohorts of Trio's daughters (Table 1), 13 cows (30%) out of his total of 44 daughters produced at least one multiple, with 21 (19%) out of 109 calvings being multiple births, and 6 sets (29%) out of 21 multiples being triplet sets. For those cows calving triplets, surprisingly two sets were produced by 2-year-olds, two sets by 3-year-olds, and the one cow with two sets calved these as a 3-year-old and a 4-year-old. 3.4. Calf survival Calf survival for the triplet-born calves from Trio's daughters was low. Eight calves (44%) survived from six sets. One of these sets was premature (all died). For the other five sets, denoting A = born alive and D = born dead, there were two AAA sets, two DDD sets and one AAD. No post mortem investigations were carried out. Adding data from the original three sets produced by Treble herself, 12 calves (also 44%) survived from nine sets. 3.5. Associated animals and data 1. Blood samples for DNA were collected in 2006 from 42 daughters of Trio, for molecular study at the University of Wisconsin, USA. DNA has also been stored from Treble herself. 2. Semen was collected from Trio in 2006, and exported to Wisconsin for one of us (BWK) to generate more daughters by artificial insemination and embryo transplant. A total of 36 Trio daughters were born in the USA in 2008 and are currently being evaluated as yearlings for ovulation rate.
C.A. Morris et al. / Livestock Science 128 (2010) 193–196
195
Table 1 Calving history up to 2008 for ‘Trio’-sired cows born at AgResearch and on the Levet property. Birth location Year of birth
Calving years
Calving location Number of cows Cows with multiples Number of calvings b Twin sets Triplet sets
AgResearch
2000 and 2001 2002 and 2003 AgResearch 2004–07 AgResearch 2004–08 Levet
Levet
2002 2003 2004
Total
2004–08 2005–08 2006–08
Levet Levet Levet
14 a (1) (10) 14 4 10 16 30 44
3a (1) (0) 3 2 4 4 10 13
18 3 35 56 10 19 24 53 109
3 1 0 4 2 5 4 11 15
1 0 0 1 1 2 2 5 6
a
The 14 cows born at AgResearch and calving at AgResearch up to 2003 had post-2003 calvings in two locations, shown in the subsequent two rows (three other cows did not calve again). b Includes multiples (combining twin and triplet sets) and single calvings.
Approximately 55 and 35 additional Trio daughters are expected to be born in 2009 and 2010, respectively. All these offspring will be used for ovulation research and molecular genetic studies. 3. Nuclear transfer was performed with skin fibroblasts obtained from an ear punch from Treble, and two clones were generated in 2000 (D.N. Wells, AgResearch, personal communication: 2000). At ∼ 18 months of age, these clones were observed with triplet and quadruplet ovulations after normal oestrous patterns, although triplet and quadruplet ovulations are otherwise a rare phenomenon in cattle. By 2007, one of the clones produced only one set of twins. 4. Discussion This report describes an elite cow, Treble, which calved three sets of triplets, and had exceptional reproductive performance in her descendants, especially six sets of triplets produced by some of her 44 grand-daughters descended through one son, Trio. The high incidence of twin- and tripletproducing animals (13 (30%) of all grand-daughters) was consistent with the high performance of some prolific cow families resulting from nearly 20 years of screening and selection for twinning in an AgResearch herd (Morris and Wheeler, 2002), but with the additional attribute of triplet calvings (six (29%) out of 21 multiple births were triplet sets). In the experimental twinning herd at Clay Center, Nebraska, Echternkamp et al. (2007a) found that cows recorded with 1, 2 or 3 foetuses had, on average, 95.7 ± 0.6%, 87.8 ± 0.8% and 54.9 ± 2.3% pregnancies maintained to term, respectively. Echternkamp et al. (2007b) suggested that triplet pregnancies in cattle were compromised in dams of all ages, by physical or physiological limitations. It is therefore suggested that cows in Treble's family may have not only high ovulation rates but also sufficient uterine capacity to carry calves to term. If this is the result of a mutation of a major gene for ovulation rate, this mutation would need to be present in a background which provides for high uterine capacity, and that background would still need to be sufficient when diluted to 25% in grand-daughters. For single-gene inheritance, a gene or gene complex would need to be transmitted as a single copy from Treble to some granddaughters via a son, and such a gene would also need to be expressed in females as a dominant or partially dominant.
The data on Trio's daughters described in Table 1 (30% producing at least one multiple, 19% of all calvings being multiple births, and 29% of the multiples being triplet sets) can be compared with: A. the experimental twinning herd at AgResearch which had relatively high twinning rates (excluding the Trio family, there were 33% twin births in six calving years, 2003–08; C.A. Morris, unpublished data: 2009) but no triplets, and B. other cows on the Levet property derived from the AgResearch high twinning rate herd, without any known ancestral link to Trio or Treble, and their figures were 14 sets of twins (12.6%) from 111 calvings, and no triplets. The cow performance details in Table 1 needed to be separated into cow-birth years because the cows had different groups of foundation dams, from the AgResearch twin-breeding herd or the unselected Levet herd. Attributing the triplet incidence to a cow-sire effect might be uncertain in the case of those derived from AgResearch dams because of their experimental selection history; however, in 26 years in the AgResearch herd, there have been no other cases of triplets, from 953 cows with 2677 calvings. Some other exceptional cows or cow families have been reported in 10 historical references reviewed by Morris (1984), including a ‘Black Polled’ cow with 25 calves from eight calvings (one single, two sets each of twins, triplets and quadruplets, and a set of sextuplets), and a cow with 15 calves from five calvings (a single and two sets each of triplets and quadruplets). In the Clay Center, Nebraska, herd above and in North American Holsteins, Kim et al. (2009) have reported successful genomic searches for quantitative trait loci (QTL) on bovine chromosome (BTA) 5 for multiple births, and a Norwegian study has shown QTL on BTA 5, 7, 12 and 23 (Lien et al., 2000). Other QTL locations have also been reported by Arias and Kirkpatrick (2004) and by Cobanoglu et al. (2005). The Treble family should provide the basis for a similar genomic study, including DNA samples from Trio's daughters in the Levet herd, New Zealand, and also from Trio's daughters born in the USA following the importation of his semen there. In conclusion, Treble is a cow of remarkable reproductive performance, with three sets of triplets. The fact that some of her grand-daughters also produced triplets strongly suggests that some of her ability is inherited. Treble and her family should provide a valuable resource for further study of variation in bovine fecundity.
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