Dairy Sheep Symposium - the Department of Animal Sciences ...

Dairy Sheep Symposium - the Department of Animal Sciences ... Dairy Sheep Symposium - the Department of Animal Sciences ...

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0.05) during the treatment and post-treatment periods, respectively, for CLY ewes compared to CLN ewes (Table 1) and peaked on d 9 for CLY ewes (Figure 2B). Average milk fat and milk protein percentage for CLY ewes was lowest (P < 0.05) during the treatment period compared to the pre-treatment period (Table 1). Average milk fat yield for CLY ewes was similar during the entire experiment (87.4 g/d), however average milk protein yield was lower (P < 0.05) during the post-treatment period (70.8 g/d) compared to the pre-treatment and treatment periods (81.7 g/d, Table 1). During the treatment period, daily milk fat and protein yield for CLY ewes tended (P < 0.10) to increase from d 5 to13, decrease (P < 0.05) from d 13 to d 18, and remain stable during the post-treatment period (Figure 3). Milk fat yield for CLN ewes tended (P < 0.10) to increase from d 5 to 7, and then decreased until d 18; milk protein yield for CLN ewes followed the same pattern, but peaked between d 6 to 8, and decreased until the end of the experiment (Figure 3). Serum P4 concentration was similar for CLY and CLN ewes from d 0 to 4 (2.2 ng/ml), corresponding to CL development (Table 1 and Figure 4). Concentration of P4 for CLY ewes peaked between d 6 and 10 (Figure 4) and was consistently greater (P < 0.0001) than for CLN ewes until the end of the treatment period (5.3 vs. 2.9 ng/ml, respectively, Table 1); P4 concentration for CLN ewes did not change over time during the treatment period (Figure 4). Concentration of P4 for CLY and CLN ewes was similar during the post-treatment period (0.8 ng/ml, Table 1) and did not differ from P4 measured on d 0 (Figure 4). Serum concentration of E2 was greatest (P < 0.05) during estrus, decreased by d 4, and was not different from 0 during the treatment and post-treatment periods (Figure 5). There were no differences in E2 concentration between CLY and CLN ewes during any period (Figure 5). Discussion This study provides the most direct evidence, to date, that the presence of CL increases milk production in dairy ewes. Two previous studies using superovulated Lacaune ewes (Labussière et al., 1993, 1996) implicated the CL in milk production and provided the rationale for the present research. Labussière et al. (1993) superovulated Lacaune ewes with various hormonal treatments and analyzed milk yield of ewes that had 0, 1, 2, 3 to 6, or > 6 CL after the hormonal treatments. There was a positive relationship between number of CL and milk yield with ewes in the group with > 6 CL having significantly greater milk yield than ewes with 0 CL. In the second study (Labussière et al., 1996), Lacaune ewes were either untreated, treated with a progestin sponge, or treated with a progestin sponge and superovulated with FSH and LH. Superovulated ewes had greater milk yield (+ 11.3%) and fat yield (+ 11.1%) than untreated or progestin-treated ewes, again implicating the presence of CL in milk production of dairy ewes. These previous studies found significant effects on milk production in the presence of a supraphysiological number of CL and the experimental designs could not preclude the possibility that numbers of preovulatory follicles and circulating estradiol concentrations were the cause of the increase in milk yield. In the present study, ewes were studied during the non-breeding season so that circulating estradiol and progesterone were related to exogenous hormonal treatments. All ewes received the same hormonal treatments except for a PGF 2α injection on d 5 to eliminate the CL in CLN ewes. In addition, number of CL (2.4 CL) in this study was similar to the number that would be expected in normally ovulating East Friesian crossbred ewes during the breeding season (average lambing rate of 220%, McKusick et al., 1999a). Within 5 d after regression of

the CL there were significant differences in milk yield between ewes that had or did not have CL. Thus, the experimental design of the present study allowed us to produce strong evidence in support of the hypothesis that the presence of a normal number of CL increases milk production; however, the mechanism(s) responsible for this increase are not conclusively demonstrated. It seems logical that the hormonal milieu produced by the CL is responsible for the increase in milk production and we postulate that an expected increase in circulating OT may be the key hormonal change that produces the observed increase in milk production. The corpus luteum has been found to contain high amounts of both mRNA and protein for OT (Jones and Flint, 1988) and circulating OT concentrations are much greater in ewes with CL than in ewes without CL (Labussière et al., 1993; Marnet et al., 1998; Schams et al., 1982). Although OT was not measured in the current study, other researchers have consistently found significantly elevated basal OT concentrations in ewes with CL as compared to ewes without CL. For example, circulating OT was extremely low during the 2 d prior to estrus (near lower limit of assay of 3 pg/ml), but increased to high concentrations (30-60 pg/ml) between d 5 and 10 after estrus (Schams et al., 1982). Similarly, superovulated ewes with > 6 CL had basal circulating OT concentrations of > 100 pg/ml as compared to 25 to 50 pg/ml in ewes with 1 CL (d 4 to 12 of the estrous cycle) and these concentrations were greater than the < 10 pg/ml concentration found in ewes with no CL (Labussière et al., 1993). Numerous studies have shown that exogenous OT treatment increases milk production in ruminants (Heap et al., 1986; Knight, 1994) and this occurs whether OT is administered before or after milking (Ballou et al., 1993). Daily treatment of Mehraban ewes (Zamiri et al., 2001) with 2 IU of OT from d 15 of lactation dramatically increased total milk yield (+ 55.5%) due to greater average daily milk yield as well as longer lactation length (175 vs. 143 d). Thus, a likely physiological model to explain our results is that the presence of CL increases circulating OT and this hormone increases milk production through possibly multiple physiological mechanisms. Oxytocin is critical for milk transfer from the alveoli to the cistern. Mice with knockout of the OT gene do not have milk ejection during suckling and pups die soon after birth unless exogenous OT is provided, whereas other aspects of reproduction and parturition appear normal in these mice (Young et al., 1996). Very low concentrations of OT are necessary to elicit milk ejection in ruminants (Gorewit et al., 1983). During machine milking of dairy ewes, plasma OT concentrations peak between 70 and 200 pg/ml and this OT is associated with ejection of milk from the alveoli to the cistern (Bruckmaier et al., 1997; Marnet et al., 1998). There is little scientific literature on the role of OT during the intermilking interval in ruminants. In the goat (Peaker and Blatchford, 1988) and ewe (McKusick et al., 2001, unpublished data), milk begins to accumulate in the cistern immediately following milking and continues to fill the cistern in a linear fashion until 16 to 24 hr, after which milk production begins to reach a plateau. Compared with the cow, the ewe and goat have relatively larger cisternal storage capacity resulting in substantial extra-alveolar storage of milk (Bruckmaier et al., 1997; Knight et al., 1994). Movement of milk from the alveoli to the cistern would result in less alveolar pressure (Labussière, 1988), lower concentrations of local feedback inhibitors of lactation (Wilde et al., 1987), and less alveolar milk fat (Levy, 1964; Williamson et al., 1995), all conditions that could increase milk secretion. In addition, there may be direct effects of OT on milk synthesis by mammary epithelial cells (Ballou et al., 1960; Benson et al., 1960; Ollivier-Bousquet and Courrier, 1976); although, it is difficult to rule out that stimulatory effects are the result of more efficient removal of residual alveolar milk by OT and therefore decreased feedback inhibition of lactation in the area

<strong>the</strong> CL <strong>the</strong>re were significant differences in milk yield between ewes that had or did not have CL.<br />

Thus, <strong>the</strong> experimental design <strong>of</strong> <strong>the</strong> present study allowed us to produce strong evidence in<br />

support <strong>of</strong> <strong>the</strong> hypo<strong>the</strong>sis that <strong>the</strong> presence <strong>of</strong> a normal number <strong>of</strong> CL increases milk production;<br />

however, <strong>the</strong> mechanism(s) responsible for this increase are not conclusively demonstrated.<br />

It seems logical that <strong>the</strong> hormonal milieu produced by <strong>the</strong> CL is responsible for <strong>the</strong> increase<br />

in milk production and we postulate that an expected increase in circulating OT may be <strong>the</strong> key<br />

hormonal change that produces <strong>the</strong> observed increase in milk production. The corpus luteum has<br />

been found to contain high amounts <strong>of</strong> both mRNA and protein for OT (Jones and Flint, 1988)<br />

and circulating OT concentrations are much greater in ewes with CL than in ewes without CL<br />

(Labussière et al., 1993; Marnet et al., 1998; Schams et al., 1982). Although OT was not measured<br />

in <strong>the</strong> current study, o<strong>the</strong>r researchers have consistently found significantly elevated basal<br />

OT concentrations in ewes with CL as compared to ewes without CL. For example, circulating<br />

OT was extremely low during <strong>the</strong> 2 d prior to estrus (near lower limit <strong>of</strong> assay <strong>of</strong> 3 pg/ml), but<br />

increased to high concentrations (30-60 pg/ml) between d 5 and 10 after estrus (Schams et al.,<br />

1982). Similarly, superovulated ewes with > 6 CL had basal circulating OT concentrations <strong>of</strong> ><br />

100 pg/ml as compared to 25 to 50 pg/ml in ewes with 1 CL (d 4 to 12 <strong>of</strong> <strong>the</strong> estrous cycle) and<br />

<strong>the</strong>se concentrations were greater than <strong>the</strong> < 10 pg/ml concentration found in ewes with no CL<br />

(Labussière et al., 1993). Numerous studies have shown that exogenous OT treatment increases<br />

milk production in ruminants (Heap et al., 1986; Knight, 1994) and this occurs whe<strong>the</strong>r OT is<br />

administered before or after milking (Ballou et al., 1993). Daily treatment <strong>of</strong> Mehraban ewes<br />

(Zamiri et al., 2001) with 2 IU <strong>of</strong> OT from d 15 <strong>of</strong> lactation dramatically increased total milk<br />

yield (+ 55.5%) due to greater average daily milk yield as well as longer lactation length (175 vs.<br />

143 d). Thus, a likely physiological model to explain our results is that <strong>the</strong> presence <strong>of</strong> CL<br />

increases circulating OT and this hormone increases milk production through possibly multiple<br />

physiological mechanisms.<br />

Oxytocin is critical for milk transfer from <strong>the</strong> alveoli to <strong>the</strong> cistern. Mice with knockout <strong>of</strong><br />

<strong>the</strong> OT gene do not have milk ejection during suckling and pups die soon after birth unless<br />

exogenous OT is provided, whereas o<strong>the</strong>r aspects <strong>of</strong> reproduction and parturition appear normal<br />

in <strong>the</strong>se mice (Young et al., 1996). Very low concentrations <strong>of</strong> OT are necessary to elicit milk<br />

ejection in ruminants (Gorewit et al., 1983). During machine milking <strong>of</strong> dairy ewes, plasma OT<br />

concentrations peak between 70 and 200 pg/ml and this OT is associated with ejection <strong>of</strong> milk<br />

from <strong>the</strong> alveoli to <strong>the</strong> cistern (Bruckmaier et al., 1997; Marnet et al., 1998). There is little<br />

scientific literature on <strong>the</strong> role <strong>of</strong> OT during <strong>the</strong> intermilking interval in ruminants. In <strong>the</strong> goat<br />

(Peaker and Blatchford, 1988) and ewe (McKusick et al., 2001, unpublished data), milk begins to<br />

accumulate in <strong>the</strong> cistern immediately following milking and continues to fill <strong>the</strong> cistern in a<br />

linear fashion until 16 to 24 hr, after which milk production begins to reach a plateau. Compared<br />

with <strong>the</strong> cow, <strong>the</strong> ewe and goat have relatively larger cisternal storage capacity resulting in<br />

substantial extra-alveolar storage <strong>of</strong> milk (Bruckmaier et al., 1997; Knight et al., 1994). Movement<br />

<strong>of</strong> milk from <strong>the</strong> alveoli to <strong>the</strong> cistern would result in less alveolar pressure (Labussière,<br />

1988), lower concentrations <strong>of</strong> local feedback inhibitors <strong>of</strong> lactation (Wilde et al., 1987), and less<br />

alveolar milk fat (Levy, 1964; Williamson et al., 1995), all conditions that could increase milk<br />

secretion. In addition, <strong>the</strong>re may be direct effects <strong>of</strong> OT on milk syn<strong>the</strong>sis by mammary epi<strong>the</strong>lial<br />

cells (Ballou et al., 1960; Benson et al., 1960; Ollivier-Bousquet and Courrier, 1976); although,<br />

it is difficult to rule out that stimulatory effects are <strong>the</strong> result <strong>of</strong> more efficient removal <strong>of</strong><br />

residual alveolar milk by OT and <strong>the</strong>refore decreased feedback inhibition <strong>of</strong> lactation in <strong>the</strong> area

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