Methods to Improve Profitability
Through Hyperprolificacy
Thomas E. Long
National
Pig Development (USA)
Introduction
Reproductive traits (e.g., number of piglets born
alive per litter and pre-weaning viability) are important components of the
economic efficiency of a swine production unit (Tess et al., 1983; de Vries,
1989a; Rothschild and Bidanel, 1999). Some swine geneticists have noted that as
genetic improvements are realized by a number of breeding organizations in the
growth and carcass leanness of their products, reproductive efficiency will
become more important as a selection objective (Ollivier et al., 1990; Webb,
1991; Schinckel et al., 1998). The magnitude of this shift in selection
objectives for dam lines of an individual breeding organization will depend on
that organization’s competitive position within the industry (de Vries, 1989b).
However, evidence of the current importance of genetic improvement in
reproductive efficiency in swine in the US can be seen in NPPC’s recent
Maternal Line Genetic Evaluation Program (2000) and by NSIF including a
symposium on hyperprolificacy at its 25th anniversary meeting. While
there are a number of traits, in addition to number born alive, that affect
reproductive efficiency in sows (e.g., piglet birth weight, piglet survivability,
farrowing interval, longevity), the focus of this paper will be on methods to
increase number born alive per litter to “hyperprolific” levels.
“Traditional” Selection
for Litter Size
Before discussing hyperprolific lines and schemes, one
point needs to be made. Number born alive does have a relatively low
heritability, and a review of studies investigating this trait found a pooled
estimate of heritability for NBA of 0.10 (Hermesch, 1996). This does not mean
that this trait cannot be substantially improved through selecting animals on a
dam index over time. A number of groups have improved this trait using
selection index and BLUP procedures in their lines (Lofgren et al., 1994, Short
et al. 1994, Long, unpublished data). BLUP applications have been implemented
in the US swine industry for genetic evaluation of reproductive traits (e.g.,
STAGES, Schinckel et al, 1986; PEST, Groeneveld et al. 1990; PIGBLUP, Long et
al., 1990). Use of these tools, in a well-designed selection program, can
improve number born alive, even though this trait is lowly heritable.
Component Trait Selection
Number born alive is a composite trait, made up of many
component traits (e.g., ovulation rate, embryonic/fetal survival, uterine
capacity). A great deal of work has been done in Nebraska to investigate
whether selection for component traits of litter size would result in greater
response than direct selection for litter size, both at the University of
Nebraska-Lincoln and the ARS-USDA facility at Clay Center (Cunningham et al.,
1979; Johnson et al., 1984; Bennett and Leymaster, 1989; Neal et al., 1989;
Lamberson et al., 1991; Leymaster and Johnson, 1994; Johnson et al., 1999).
Since one of the speakers at this symposium will be covering this area, it won’t
be addressed directly here. However, results from the NPPC Maternal Line
Genetic Evaluation Program (2000) would suggest that component trait selection
has been successful in developing a hyperprolific dam line, the Nebraska Index
Line.
Molecular Genetics
Approaches
Given the low heritability of litter size in swine, the
discovery of the estrogen receptor gene (Rothschild et al., 1996) and its
effects on litter size were of great interest to swine breeders everywhere.
Haley (1999) noted that the association between the ESR locus and litter size
“has been identified in a number of populations, although the magnitude of the
association varies between populations.” Since then other researchers have
reported quantitative trait loci having affects on reproductive traits (Rathje
et al., 1997; Southwood et al., 1998; Spiegel et al., 1999; Cassady et al.,
1999). Use of genetic markers in developing hyperprolific dam lines in swine
will become an important tool for swine breeders to use in the future. Use of these
tools, in practical breeding programs, will depend on issues, such as: the
availability of markers, the amount of variation a specific marker is
accounting for, combining marker information with current quantitative genetic
evaluation procedures, other genes in the genome that may have epistatic
effects with the specific allele in question and the frequency of the allele of
interest in the population being screened. As work continues in the molecular
area, there may be many more opportunities to exploit this technology to
enhance reproductive performance in dam lines.
Hyperprolific Schemes
Hyperprolific schemes have been described (Legault and
Gruand, 1976; Bichard and Seidel, 1982) and involve screening multiplier units
for hyperprolific sows such that their germplasm can be brought back into the
nucleus selection program. These types of breeding schemes do afford higher
levels of selection intensity than could be applied in the nucleus, but have
generally not been adopted due to genetic lag in production traits that would
be incurred in the process (Avalos and Smith, 1986). Bidanel et al. (1994), in
reviewing 20 years of a hyperprolific scheme in France, noted that, while
litter size was substantially improved and growth rate unaffected, these dam lines
had higher backfat and poorer feed conversion than contemporary lines.
Additionally, there are health risks that might be incurred in moving germplasm
from multiplier units, as multiplier units tend to have poorer health levels
than nucleus farms. However, as technology and swine populations change, should
hyperprolific schemes be reconsidered? Information technology has improved such
that estimated breeding values can be calculated on multiplier animals rather
than relying on phenotypic data. As stated previously, many organizations have
realized genetic improvements in dam lines for production traits, such that
those organizations have favorable competitive positions for those traits.
Could hyperprolific schemes be implemented in these systems, such that there
was an enhancement of reproductive performance with smaller penalties for
genetic lag in production traits than would have been incurred in the past?
Finally, embryo transfer protocols may afford opportunities to move germplasm
from multiplication hyperprolific sows to nucleus units in ways that minimize
health risks. Although hyperprolific breeding schemes are probably not being
used in most current dam line breeding programs, in a discussion of the
development and use of hyperprolific lines, one should raise the question of
whether these types of schemes should be revisited.
Use of Hyperprolific
Lines
This paper has discussed methods of developing
hyperprolific lines. Once these lines have been developed within a breeding
organization or samples (animals, semen or embryos) from a hyperprolific line
have been obtained, it needs to be determined how those lines will be used in
breeding programs in combination with other dam and sire lines to produce
market pigs, since many hyperprolific dam lines tend to have poorer growth and
carcass characteristics than more “standard” dam lines (Bidanel, 1990; Bidanel
et al., 1994; NPPC, 2000). The two obvious alternatives are: 1) use the line in
a specific crossing program with other dam lines and superior terminal sire
lines, or 2) develop new composite lines using the hyperprolific line as one of
the foundation lines. Both these approaches have or are being implemented by
breeding organizations. Using the hyperprolific line in a specific crossing
program is relatively straightforward, but does rely on having superior
terminal lines to use in breed combinations. Results from the NPPC Terminal
Sire Line Genetic Evaluation Program (1995) and the NPPC Lean Growth Modeling
Project (1998) clearly indicate that there are terminal sire line differences
for production and carcass characteristics. Therefore, decisions on what type
of terminal sire line to use in combination with hyperprolific lines are
important for production and processing sectors to benefit from the use of
these lines.
Examples of the development of hyperprolific
composite lines can be found in breeding programs using Chinese breeds, such as
the Meishan or Jiaxing, in combination with Western dam lines (Mercer and
Hoste, 1994; Burlot et al., 1998). These programs depend on also using Western
dam lines, as foundation lines, that are not only prolific, but also have
excellent production characteristics. Selection objectives in these composite
lines are distinct from usual dam line breeding objectives being applied in
most current breeding programs, and the use of these composite lines in
crossing programs also relies on superior terminal sire lines to produce market
pigs.
The importance of reproduction to the economic
efficiency of swine production has been previously stated. Trends over the last
5-7 years from USDA Pig Crop reports indicate an increase in the number of pigs
being produced per sow. Industry analysts have provided a number of reasons for
these trends, such as changing demographics of swine production, improved
husbandry practices and better genetic lines. It is clear that what were once
good reproductive output levels from sows may now only be ordinary. The use of
hyperprolific dam lines in a breeding program is one way to improve reproductive
output from a swine operation.
References
Avalos, E. and C. Smith. 1987. Genetic improvement
of litter size in pigs. Anim. Prod.
44:153-164.
Bennett,G.L. and K.A.
Leymaster.
1989. Integration of ovulation rate, potential embryonic viability and uterine
capacity into a model of litter size in swine. J. Anim. Sci. 67:1230-1241.
Bichard, M. and C.M. Seidel. 1982. Selection for
reproduction performance in maternal lines of pigs. Proc 2nd World
Cong. on Genet. Appl. to Livestk. Prod.
Bidanel, J.P. 1990. Potential use of
prolific Chinese breeds in maternal lines of pigs. Proc. 4th World
Cong. on Genet. Appl. to Livestk. Prod. XV:481-484.
Bidanel, J.P., J. Gruand and
C. Legault.
1994. An overview of twenty years of selection for litter size in pigs using
‘hyperprolific’ schemes. Proc. 5th World Cong. on Genet. Appl. to
Livestk. Prod. 17:512-515.
Burlot, T., Z. Siqing, J.
Naveau, C. Legault and J.P. Bidanel. 1998. Genetic parameters and genetic trends in the
Sin-European Tiameslan composite line. Proc. 6th World Cong. Genet.
Appl. to Livestk. Prod. 23:599-602.
Cassady, J.P., R.K. Johnson,
D. Pomp, L.D. Van Vleck, E.K. Speigel, K.M. Gilson and G.A. Rohrer. 1999. Evidence for QTL on
chromosome 1, 8 and 13 affecting reproduction in pigs. J. Anim. Sci. 77:
Suppl.1:3.
Cunningham,P.J., M.E.
England, L.D. Young and D.R. Zimmerman. 1979. Selection for ovulation rate in swine:
Correlated response in litter size and weight. J. Anim. Sci. 48:509-516.
de Vries, A. G. 1989a. A model to estimate
economic value of traits in pig breeding. Livestk. Prod. Sci. 21:49-66.
de Vries, A. G. 1989b. A method to
incorporate competitive position in the breeding goal. Anim. Prod. 48:221-227.
Groeneveld, E., M. Kovac, T.
Wang. 1990.
PEST, a general purpose BLUP package for multivariate prediction and
estimation. Proc. of 4th World Cong. on Genet. Appl. To Livestk.
Prod. Edinburgh. XIII:488-491.
Haley, C. 1999. Advances in
quantitative trait locus mapping. In “From Jay Lush to Genomics: Visions for
Animal Breeding and Genetics”. Eds. J.C.M. Dekkers, S.J. Lamont and M.F.
Rothschild. Iowa State University, Ames.
Hermesch, S. 1996. Genetic parameters
for lean meat yield, meat quality, reproduction and feed efficiency traits for
Australian pigs. PhD Dissertation. AGBU, UNE, Armidale, NSW, Australia.
Johnson, R.K., D.R.
Zimmerman and R.J. Kittok. 1984. Selection for components of reproduction in swine. Livestk.
Prod. Sci. 11:541-558.
Johnson, R.K., M.K. Nielsen
and D.S. Casey.
1999. Responses in ovulation rate, embryonal survival and litter traits in
swine to 14 generations of selection to increase litter size. J. Anim. Sci.
77:541-557.
Lamberson, W.R., R.K.
Johnson, D.R. Zimmerman and T.E. Long, 1991. Direct responses to selection for increased
litter size, decreased age at puberty or random selection following selection
for ovulation rate in swine. J. Anim. Sci. 69:3129-3143.
Legault, C. and J. Gruand. 1976. Improvement of
litter size in sows by creation of a ‘hyperprolific’ line and use of artificial
insemination: theory and preliminary results. Journees Recherche Porcine
France. 8:201-206.
Leymaster, K.A. and R.K.
Johnson.
1994. Second thoughts on selection for components of reproduction in swine.
Proc 5th World Cong. on Genet. Appl. to Livestk. Prod. 17:307-314.
Lofgren, D.L., D.L. Harris,
T.S. Stewart, D.D. Anderson, A.P. Schinckel and M.E. Einstein. 1994. Genetic progress of
the US Yorkshire breed. Proc. of 5th World Cong. on Genet. Appl. To
Livestk. Prod. Guelph. 17:425-428.
Long, T., H. Brandt and K.
Hammond.
1990. Breeding value prediction with the animal model for pigs. Proc. of 4th
World Cong. on Genet. Appl. To Livestk. Prod. Edinburgh, XV:465-468.
Mercer, J.T. and S. Hoste. 1994. Prospects for the
commercial use of Chinese pigs. 17:327-334.
Neal, S.M., R.K. Johnson and
R.J. Kittok.
1989. Index selection for components of litter size in swine: Response to five
generations of selection. J. Anim. Sci. 67:1933-1945.
Ollivier, L., R. Gueblez,
A.J. Webb, and H.A.M. van der Steen. 1990. Breeding goals for nationally and internationally
operating pig breeding organizations. Proc. of 4th World Cong. on
Genet. Appl. To Livestk. Prod. Edinburgh. XV:383-394.
Rathje, T.A., G.A. Rohrer
and R.K. Johnson. 1997. Evidence for quantitative trait loci affecting ovulation rate
in pigs. J. Anim. Sci. 75: 1486-1494.
Rothschild, M.F., C.
Jacobson, D.A. Vaske, C.K. Tuggle, L. Wang, T. Short, G.R. Eckardt, S. Sasaki,
A. Vincent, D. McLaren, O. Southwood, H. van der Steen, A. Mileham and G.
Plastow.
1996. The estrogen receptor locus is associated with a major gene influencing
litter size in pigs. Proc. Natl. Acad. Sci. USA. 93:201-205.
Rothschild, M.F. and J.P.
Bidanel.
1998. Biology and genetics of reproduction. In “ The Genetics of the Pig”. Eds.
M.F. Rothschild and A. Ruvinsky. CAB International. New York, New York.
Schinckel, A.P., D.L.
Harris, T.S. Stewart and D.L. Lofgren. 1986. Swine testing and genetic evaluation system
for the purbred swine associations. Proc. of 3rd World Cong. on
Genet. Appl. to Livestk. Prod. Lincoln. X:98-109.
Schinckel, A.P., D.L.
Lofgren and M.E. Einstein. 1998. Recent STAGES index changes. Seedstock Edge, July Herdsire
Issue: 120-121.
Short, T.H., E.R. Wilson,
D.G. McLaren.
1994. Relationships between growth and litter traits in pig dam lines. Proc. of
5th World Cong. on Genet. Appl. to Livestk. Prod. Guelph.
17:413-416.
Southwood, O.I., T.H. Short,
and G.S. Plastow. 1998. Genetic markers for litter size in commercial lines of pigs.
Proc. 5th World Cong. on Genet. Appl. to Livestk. Prod. 26:453-456.
Spiegel, E.K., J.P. Cassady,
K. Gilson, R.K. Johnson, D. Pomp, L.D. Van Vleck and G. Rohrer. 1999. Indication of
quantitative trait loci on chromosome 6 affecting porcine reproductive traits.
J. Anim. Sci. 77: Suppl. 1:2.