Multi-stage selection of sires

The multi-stage selection schemes resulted in equal or lower total economic response as compared with the equivalent one-stage selection schemes (V). The relative proportions of response from different trait groups in multi-stage selection schemes were slightly different from those of one-stage selection. The asymptotic response in milk traits was somewhat re- duced, while the response in carcass traits of bulls was, in most cases, either equal or slightly higher than the response obtained with one-stage selection. All studied multi-stage selec- tion schemes resulted in an unfavourable response in carcass traits of cows due to decreased carcass fleshiness and, in some cases, also increased carcass weight.

Overall, the best multi-stage selection scheme was a three-stage scheme where 10% of sires were culled at the age of 1.5 years based on their own carcass traits. Further, 22% of sires were culled at the age of 3.5 years based on the previously available information plus information on their sons’ carcass traits. The final culling of sires was done at the age of 5.5 years based on all previously available information plus information on their daughters’ carcass and milk traits. At this final stage, the top 3% of sire candidates were selected. The scheme resulted in approximately equal total economic response to the one-stage scheme that utilized the same information, but its costs are expected to be somewhat lower due to need to maintain fewer bulls. There is still room for further optimisation of the selected proportion in different stages and selection schemes.

6

CONCLUSIONS

Based on the analyses of selection strategies, it is feasible to expand the total merit index of Finnish dairy cattle breeds to include carcass traits (IV, V). The inclusion of carcass traits in the breeding goal for both bulls and cows does not hinder progress in milk traits. It would also reflect the income sources of the farms better. The carcass weight and carcass quality traits of bulls in particular would benefit from being included in the breeding goal.

Results show that it is possible to breed for improved carcass weight and quality of young FAy and HFr slaughter animals. The carcass traits are heritable, although the heritability for the bull and heifer traits is relatively low (I). However, heritability of carcass traits of bulls and heifers is higher than, for example, the fertility and health traits that are currently included in breeding goals of dairy cattle. The genetic correlations of bull and heifer carcass traits are generally favourable, both between the carcass traits themselves (I) and between the carcass traits and milk production traits (III).

The relatively high heritability of carcass traits of FAy cows suggest that it would be easy to breed for improved carcass quality and optimal body weight of dairy cows (II). Although dairy cows are not intended as beef-producing animals, improved carcass fatness and flesh- iness, and thus, improved body condition might help to reduce stress caused by lactation in high-yielding cows. This could, in turn, reduce problems related to negative energy balance during early lactation, such as weakened fertility.

Due to the negative genetic correlation between carcass quality and milk traits of dairy cows, their fatness and fleshiness will continue to deteriorate as a result of selection on milk production traits unless some counteractive selection is practised (II, V). The high positive genetic correlation between carcass traits of bulls/heifers and carcass traits of dairy cows might help to reduce the deterioration of cow carcass quality if improved carcass traits of young slaughter animals are selected for (III, V).

The data for progeny testing young AI-sires for carcass traits is already available due to the evaluation system of AI-sires in Finland. The progeny carcass data can be collected at a fraction of the cost of the data needed for progeny testing of sires for milk production traits. Furthermore, simplified evaluations that ignore genetic and phenotypic correlations between carcass and milk traits can be used for the prediction of breeding values of dairy sires without fear of reduced total selection response (IV).

Because bull and heifer progeny of young sires are slaughtered approximately 1.5 years before sires’ daughters finish their first lactation, it would be possible to select young sires on carcass traits over a year earlier than for milk. According to the results of this study, however, selection of sires for their carcass and milk traits could be done in one stage at the age of 5.5 years, since multi-stage selection of sires does not yield extra profit over one-stage selection with respect to the total selection response (V). It should be noted that in this study the selection intensity of sires was fixed at each stage, and thus, was not necessarily optimal. In future, the intensities should be optimized between the stages to maximize total selection response. Furthermore, the costs of maintaining the candidate sires were not considered in this study.

Reliable EBVs for carcass traits can be estimated for young dairy AI-sires based solely on their male progeny, which form the majority of the carcass data (V). For most sires, some daughters will also be slaughtered before they begin lactation. Including the carcass traits of these heifers in breeding evaluation might be of value in predicting fatness more accurately, as heifers provide additional information on variation in fatness, while most bulls classify in

fatness grade 2 (I, Figure 3). Combining bull and heifer data might also reduce the number of herds that provide only few carcasses, thereby improving the overall structure of data (I).

If body weight and condition of dairy cows are of interest, including some cow carcass data in selection decisions is necessary (V). Collecting carcass data on mature half-sisters and/or daughters of AI-sires would offer an easy and inexpensive way to obtain objective information on these traits. According to data of this study, approximately 10% of dairy cows were culled after the first lactation, and altogether one-third of dairy cows were culled before the third lactation. The culled half-sisters and possibly also daughters would enable an earlier genetic evaluation of cow carcass traits for the proven bulls that continue in AI-use for a few years.

Young dairy breed test bulls intended as AI-sires still go through an individual growth test in Finland, and the records from the test station are used to predict the EBV for growth. The growth EBV was omitted from the total merit index in 1993, and its importance in selection of dairy breed sires has since declined. In future, if all dairy sires get progeny test result for carcass weight and carcass quality traits, the need for an individual growth test will have to be reconsidered. Even assuming a genetic correlation of 1 between the sire’s own growth and the various carcass traits of his progeny, the economic response in carcass traits to selection on own growth is vastly inferior to the selection on progeny carcass traits (V).

Large differences are present between proven sires with respect to carcass EBVs. Particu- larly amongst the FAy, individual sires can be found that combine excellent milk production, fertility and health traits with good carcass weight and carcass quality traits. Among the HFr, the situation is less promising, probably due to the rapid Holsteinization of the breed in recent years.

The breeding goal of dairy cattle breeds should reflect the economically important traits, i.e., both beef and milk traits. If animal breeding associations do not wish to combine the beef and milk traits in one index, they could promote the use of sires with good carcass EBVs by, for example, annually naming some "dual-purpose" sires with both promising milk and beef traits. However, breeding values of the sires with respect to calving difficulties and calf mortality should also be taken into account.

7

ACKNOWLEDGEMENTS

This study was carried out at the Department of Animal Science of the University of Hel- sinki, and at the Department of Animal Sciences of the Wageningen University and Research Centre in The Netherlands. I thank both institutes for providing their facilities, and the staff for their kind assistance. It has been a pleasure to work at both places!

I want to thank specifically several people who have contributed to this work in one way or another:

• First of all I have to thank my two supervisors, Professor Matti Ojala (Department of Animal Science, University of Helsinki) and Professor Johan van Arendonk (De- partment of Animal Sciences, Wageningen University and Research Centre). Matti has always provided me with a freedom to do my own things and try my own views, whereas Johan quite literally opened up the world for me and keeps pointing out that there is life outside the borders of Finland too. It has been sometimes quite challenging when dealing with these two gentlemen, but that is what keeps things interesting! • I would also like to thank my other co-authors Päivi Parkkonen (Animal Production

Research, Agricultural Research Centre of Finland) and Dr. Piter Bijma and Marc Rut- ten (Department of Animal Sciences, Wageningen University and Research Centre). Päivi did an enormous amount of work for the Paper I, and in addition wrote a very nice manuscript of it. Piter and Marc wrote the program I used in Paper V, and it is only due to them that I was able to write the Paper V in the first place.

• Professor Johann Sölkner (Department of Livestock Sciences, University of Agricul- tural Sciences Vienna, Austria) is warmly thanked for agreeing to be my opponent. • Professor Asko Mäki-Tanila (Animal Production Research, Agricultural Research

Centre of Finland) and Dr. Erling Strandberg (Department of Animal Breeding and Genetics, Swedish University of Agricultural Sciences, Sweden) are warmly thanked for pre-examining my thesis in a record time, and for offering valuable suggestions to further improve the manuscript.

• Tarja Korhonen (Finnish Animal Breeding Association) had a crucial role in the begin- nings of the project that would later form the core of my thesis. In addition, she has offered valuable advice through the years and kindly checked through the manuscript of my thesis.

• Veijo Vilva (Department of Animal Science, University of Helsinki) has been an irre- placeable person as far as computers are concerned. I owe to him my deepest gratitude for rescuing me from numerous small and larger crises related to problematic programs and computers.

• I thank my former and present colleagues at the Department of Animal Science, Uni- versity of Helsinki for sharing the everyday life during these past years. In particular, I want to thank Minna Toivonen, Tiina Ikonen, Outi Ruottinen and Katariina Mäki for friendship, encouragement and many laughs we have shared both in our animal breed- ers’ attic and around the world on various congress and course trips. The same thanks apply also to the folks at the Department of Animal Sciences, Wageningen University and Research Centre, especially to Liesbeth van der Waaij.

• Lihakunta Oy provided the field carcass data, and Tapani Hellman (Agricultural Data Processing Centre) provided several pedigree and other data sets. My warm thanks are due to both.

• Carol Ann Pelli (Language Centre, University of Helsinki) is thanked for the thorough language review of this thesis.

• This study was supported by the Ministry of Agriculture and Forestry, the Faculty of Agriculture and Forestry at the University of Helsinki, the Wageningen University and Research Centre, and the Finnish Academy. They all are gratefully acknowledged. Finally, I would like to thank my family: two-legged, four-legged and six-finned alike. My work would never have been possible without the constant support of my parents, both men- tally and also in taking care of my zoo whenever needed. The two Departmental Dogs Pumi and Buster brought joy to all my working days, dragged me out for walks afterwards no mat- ter what weather, and in general saw after that I didn’t totally mummify behind my desk. The various fish taught me valuable lessons about breeding in practice, and in case of some very special goldfish also provided me with company during my travels through Europe. And last but certainly not least, my whole journey into the wonderful world of animals began in 1976 with the arrival of Alpertti the tortoise, whose presence continues very strongly now 24 years after and who always tries to teach me to festina lente.

8

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