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Periparturient changes in mammary gland secretions in multiparous buffalo cows (Bubalus bubalis).

INTRODUCTION

Buffalo is considered to be one of the important dairy animals in a number of Mediterranean and SouthEast Asian countries. In Africa, the census of buffalo population is estimated to be 2.2% of total buffalo around the world which is presented in Egypt. The Egyptian buffalo population is 4,200,000 head with a total annual fresh milk production of 2,614,500 tones, representing ~45% of total milk production [1].

Buffalo milk is characterized by high levels of fat, lactose, protein, ash, [Ca.sup.++], vitamins A and C and lower levels of vitamin E, riboflavin and cholesterol when compared to Holstein cow's milk [2], [3]. Moreover, the fatty acid analysis revealed that saturated fatty acids (mainly palmitic acid), trans fatty acids, linolenic acid and conjugated linolenic acid were higher in buffalo milk than in cow milk [4]. These facts give a relatively important advantage of buffalo's milk over cow's milk.

Colostrum is a unique mammary secretory product that is formed during late pregnancy when mammary cells are proliferating and differentiating for the preparation of lactation [5]. Moreover, it is a complex biological fluid, which helps in the development of newborn immunity and it contains significant quantities of complement components that act as natural anti-microbial agents to actively stimulate the maturation of an infant's immune system [6].

Colostrogenesis, identified as the prepartum transfer and accumulation of immunoglobulins from maternal circulation into mammary secretions, is a discrete and finite stage of mammogenesis. In domestic ruminants, the transfer begins several weeks prior to parturition and ceases abruptly immediately prior to parturition [7].

Many reports have identified the prepartum bovine mammary gland secretion as honey-like, which is high in protein and low in lactose. McDouoall [8] described bovine precolostrum as "a viscous honey-like substance obtained from the udders of pregnant heifers at half term". Hafez [9], also stated that early bovine prepartum secretions starting -45 days prepartum which had the consistency of honey with quantities generally small. Mammary secretions begin to accumulate soon after the formation of alveoli during pregnancy [10]. Colostrogenesis in the bovine species, the process of forming colostrum, occurs 3-5 weeks prior to parturition [7]. Moreover, offspring of bovine species are born agammaglobulinemic and immunoglobulin transmission occurs via mammary secretions (ungulates such as horses, pigs and goats) [11]. This fact gives more importance of the quality and quantity of the produced colostrum which is relatively crucial for the survival of newborn calves.

Up to the knowledge of the authors, lactogenesis and milk synthesis of buffalo cows is poorly characterized and not well documented. Therefore, the aim of the present study was to characterize the changes in the gross chemical composition of the precolostrum and colostrum with reference to their gradual changes relative to parturition.

MATERIALS AND METHODS

Animals and management:

A total of ten healthy multiparous late pregnant buffalo cows in their third to ninth parity with mean body weight of 725 [+ or -] 8 kg were recruited from the Agriculture Research Station, Faculty of Agriculture, Cairo University, Giza, Egypt. The experiment started 30 days before the expected calving date until the fourth day of lactation. Animals were housed in semi-shaded open yards and were given their nutritional allowances according to El-Ashrey et al. [12].

Mammary secretions samples and analysis:

All animals were hand milked twice a week after a proper stimulation of the udder before the morning feeding starting 30 days before the expected calving date. Starting from parturition animals were machine milked at 07:00 a.m. and 07:00 p.m. daily. Mammary secretions yields were recorded at each milking pre- and post-partum to the nearest 1 g. Precolostrm and colostrum samples were collected, at each milking in a 50 ml clean tube, and stored at -20[degrees]C for further analysis. Fat, protein, lactose, ash, total solids and solid not fat % of precolostrum and colostrum samples were measured using infrared milk analyzer instrument (Bentley 150[TM], Bentley Instruments, Inc., Chaska, Minnesota, USA). Electrical resistance of precolostrum and colostrum samples was measured using hand-held device, (Mastitis Detector, Draminski Electronics in Agriculture, Poland) according to the manufacturer the lower the device reading, the higher the likelihood of infection.

Blood samples and analysis:

Peripheral blood samples were weekly withdrawn throughout the prepartum period, on the day of parturition and on the 7th day postpartum via jugular venipuncture in a heparinized tube after morning milking and before feeding except for the samples at the day of parturition, which were taken after milking of the 1st colostrum. The blood plasma was separated immediately, by centrifugation at 5000 rpm for 15 min. at 4[degrees]C, and stored at -20[degrees]C pending hormonal analysis. Blood plasma progesterone levels were measured using ELISA kit (BioCheck, Inc., California, USA), using the ELISA reader of Biotek (Bitek[TM], ELx 808). According to the manufacturer, inter and intra assay coefficient of variability were 6.3 and 3.85%, respectively. The sensitivity of the assay was 0.0625 ng/ml. The standard curve of the progesterone was from 0 to 50 ng/ml.

Statistical analysis:

Data were statistically analyzed to measure the composition changes of buffalo precolostrum and colostrum secretions over time. Data were subjected to the analysis of variance as repeated measurements (split plot in time) according to Neter et al. [13] using the general liner model of SAS [14]. Differences among means were evaluated using Duncan's new Multiple Range Test [15]. The statistical model was:

[Y.sub.ij] = [mu] + [ST.sub.i] + [E.sub.ij].

Whereas:

[Y.sub.ij] = the observation ij.

[mu] = the overall mean.

[ST.sub.i] = the effect due to the secretion type i, whereas i= 1 for precolostrum and i= 2 for colostrum or is the effect due to the time when secretion collected i, whereas i= 10 periods for precolostrum and i= 8 periods for colostrum.

[E.sub.ij] = the experimental error associated with [Y.sub.ij] observation, assumed to be randomly distributed (0, [[sigma].sup.2]).

RESULTS AND DISCUSSION

All the experimental buffalo cows calved in fall and winter months from September, 2012 to February, 2013. The actual days of prepartum experimental period was 33.3 [+ or -] 10.73 days due to difference between the expected and the actual calving date. The majority of buffalo cows did not stand for milking prepartum. During the process of prepartum milking, the animals kept stomping (a raising and lowering of the foot in one), kicking (a forward or sideways movement of the leg, without dislodging the claws place place) and kicking off claw (a kick that causes the claws to fully or partially dislodge). Prepartum milking of the buffalo involved conditioning followed by active milking of the stored prepartum secretions then stripping of the available secretions.

The changes in the gross chemical composition relative to parturition (i.e. fat, protein, lactose, ash, total solids and solids not fat), yield and electrical resistance of precolostrum and colostrum throughout the experimental period are presented in Fig (1). Precolostrum mean percentages of fat, lactose and ash were lower while protein percentage was higher (P<0.0001) than colostrum being 2.53 vs. 7.90 (-212%), 2.61 vs. 4.18 (-60%), 1.33 vs. 1.46 (-10%) and 10.97 vs. 6.44% (+41%), respectively (Table 1). The maximum amount of precolostrum collected at a given milking was 770 g/milking and occasionally no secretion was obtained. One of the experimental buffalo did not produce enough precolostrum for analysis during the entire prepartum period starting 44 days before calving. This indicates that buffalo cows did not respond to prepartum milking as what has been reported in dairy cattle but with a same variation.

In the present study, there was a gradual increase in secretion collected prepartum as parturition approaches (Fig. 1) with the highest quantity collected at 3 to 1 day preparutm (P<0.05). Greene et al. [16] reported that on the day prior to calving, 36, 33, and 31% of the experimental cows milked prepartum produced less than 4.5 kg, 4.5 to 9 kg, and>9 kg of milk, respectively. In another study, four cows out of seven produced >4 L/d of milk by 4 days prepartum [17]. Keller et al. [18] reported that the daily yield of prepartum secretions of cows milked twice a day starting 6.8 [+ or -] 0.8 days prepartum, averaged 10 [+ or -] 2 kg one day before calving with large variations among individuals ranged from 4 to 17 kg. Grummer et al. [19] also reported large variations in prepartum secretion of cows milked at day 7 to 1 prepartum that ranged from 2.5 to 10 kg/day. On the day prior to calving, two cows produced <4.5, five produced 4.5 to 9.0, and three produced >9.0 kg of milk/day. Moreover, Daniels et al. [20] stated that udder secretions of prepartum milked heifers were measurable beginning approximately 15 day before actual calving, and gradually increased to reach 16.5 kg/day at the day of calving.

The low yield of precolostrum collected from buffalo may be due to the failure of stimulating buffalo udder epithelial cells to synthe=sis milk prepartum under the influence of high preparutm blood progesterone level (Fig. 2), which serve as a suppressor of the periparturient synthesis of milk components (e.g. a-lactalbumin and casein) [21], [22], [23]. Moreover, the process of colostrogenesis has been shown to be initiated under the influence of hormones and occurs during the last weeks of pregnancy when steroid concentrations of estradiol (E2) and progesterone (P4) are highly elevated [7], [24], [25]. Another explanation could be the low proportion of cisternal milk reported in buffalo, 95% of the milk is stored in the secretory tissue even after a milking interval of 10 to 12 hours that can only be extract with an active milk ejection [26]. Nevertheless, in the present study, yield of the first collected colostrum was the highest compared to subsequent colostrum milking (P<0.05) (Fig 1), which is in accordance with the same trend observed at calving of prepartum milked primiparous cows [27].

Chemical composition of prepartum mammary secretions is presented in Table (2). Precolostrum obtained prepartum can be characterized as sticky pale yellow honey-like secretion which dries fast when exposed to air. As parturition approached precolostrum turned to be as colostrum in appearance. Mean precolostum fat% was very low compared to its percentage in colostrum (2.53 vs. 7.90%). This result can be explained by what Akers et al. [28] have reported that there was a highly significant increase in the capacity to synthesize fatty acids from acetate, which was 6.3-fold greater, in mammary tissue from postpartum cows than from prepartum cows.

Colostrum chemical composition is presented in Table (3). Colostrum fat% postpartum started with low concentration (6.97%) which increased significantly in the second milking after 12 hrs. (10.28%) and then gradual decrease as lactation advanced. Conte and Scarantino [29] reported a same trend that the first bovine colostrum fat% was 5.10% increased to be 6.85% in the second milking 6 hours postpartum and afterword continued to decrease to reach 2.8%, 48 hours postpartum.

Lactose has been detected 34 to 32 days prepartum (1.96%) in precolostrum secretion, although being significantly lower (P<0.05) than samples from 4 to 7 days postpartum (2.84%). Another increase was observed in 1 to 3 days prepartum (3.63%) being significantly higher (P<0.05) than all the previous prepartum samples. Convey [21] referred that prepartum appearance of lactose in the mammary gland may be considered as "lactogenesis" which mean the onset of milk production. Lactose% was relatively low at all prepartum milkings except the one before parturition, which was significantly higher than the previous milkings, being close to the value of the first postpartum colostrum collected postpartum, 3.63 vs. 3.81%, respectively. This result is in consistent with what Convey [21] has reviewed that lactose in milk from cows subjected to prepartum milking increased from 10 days prepartum through parturition. The explanation of these increases in lactose% could be the increasing activity of enzymes related to lactose synthesis which appear in bovine mammary tissue by 30 days before parturition with increasing activity of 1.5 to 2.5 times from late gestation to early lactation [21]. Colostrum lactose% showed a gradual increase starting the day of parturition until reached its higher concentration 4.74% in day 4 postpartum.

There was no observed statistical difference among precolostrum protein percentages from samples collected 34 to 4 days prepartum, whereas, the samples collected 1 to 3 day prepartum were significantly lower than some earlier samples. Meanwhile, a gradual decrease was observed in precolostum protein% until reached its lowest value 10.07% at -1 to -3 days before calving, while in the first postpartum milking was 10.14%. A gradual decrease was observed in colostrum protein% starting first postpartum milking until reached the lowest value (P<0.05) at 84 hours postpartum (from 10.14 to 5.00%). The high protein% in both precolostum and first colostrum secretions shown in the present study could be explained with the specific transfer of IgG1 from the blood to mammary secretions which distinguish colostrogenesis stage in bovine [24]. Moreover, Singh et al.

[30] reported a decrease in the concentrations of buffalo colostral Igs from 10.3 to 1.4% in the first 70 hours from parturition. This transfer of immunoglobulins is reflected in the high total protein% observed in both pre- and early post-partum secretions. Dang et al. [31] stated that levels of IgG averaged 54.0 mg/ml at calving in lactating Murrah buffalo, and then decreased significantly during the five days postpartum.

The current results are in consistent with those reported by Coroian et al. [32] that buffalo colostrum fat% for the first 4 days postpartum showed mean values of 11.31 to 8.14% in summer season and 11.22 to 8.11% in winter season. Meanwhile, these values decreased gradually starting with first day postpartum until day seven postpartum. Protein and total solids showed a gradual decrease from parturition to day 4 postpartum (8.69 to 5.81% and 25.28 to 19.81%, respectively) [32]. On the other hand, lactose had an opposite trend being low at the day of parturition (3.72%) and gradually increased to reach the normal value in milk (4.95%) at day 4 postpartum. Although colostrum protein% reported by [33], [34] was relatively higher in the first milking postpartum than what we find, the protein% followed the same trend as they reported being high on the day of parturition and gradually decrease to reach ~6.5% at the fourth day postpartum and ~5.00% at 96 hours postpartum. El-Loly and Mansour [35] reported that the total solids, protein, fat and ash contents of Egyptian buffalo colostrum were higher after parturition from 0 to 12 hours postpartum being 30.40, 18.44, 7.32 and 1.07%, respectively, and then decreased rapidly to reach values of normal milk almost after three days of parturition being 15.15, 4.18, 4.7 and 0.79%, respectively. Their result showed that the colostrum protein content was the most variable constituent (18.44 vs. 4.18%). Meanwhile, the changes in the lactose content followed an opposite trend (3.27 vs. 5.44%). Abd El-Fattah et al. [34] stated that both buffalo and cow colostrum composition were approached to the normal milk composition during the first five days after parturition.

In the present study, the mean precolostrum ash% was 1.33% while in colostrum it was 1.46%. The high ash% in pre- and early post-partum milking is due to the high mineral content reported in buffalo milk [2]. It has been reported that ash started with 0.93% at the day of parturition and increase at day 6 and 7 postpartum to reach 1.02% [32].

Electrical conductivity (EC) is a measure of the resistance of milk to an electric current, conductivity is the reciprocal of the resistance. In milk, EC is determined by the concentration of anions and cations, primarily [Na.sup.+], [K.sup.+] and [Cl.sup.-]. Typical EC of milk from an uninfected cow varies between 4.0 and 5.5 mS/cm at 25[degrees]C. The present prepartum secretion electrical resistance (ER) was low compared to colostrum ER 349 vs. 445 units of electrical resistance. The ER of the first two milkings prepartum was significantly lower than the one before parturition 317, 325 and 393 units, respectively. Meanwhile, the mean postpartum ER was 445 units with no differences between milkings. The observed difference between pre- and postpartum ER values can be attributed to the tight junction permeability that increases the passage of ions and cations before closure during the onset of copious milk secretion [36]. The concept behind reading EC or ER for milk to determine mastatic milk is that destruction of tight junctions, during infection with mastitis, causes a decrease in milk lactose and [K.sup.+] concentration and increase in [Na.sup.+] and [Cl.sup.-] concentrations as a cause of the increased blood capillary permeability and the destruction of action ion-pumping systems [37], [38]. Another explanation would be the high fat%, which is a poor electrical conductor, found in colostrum compared to precolostrum (7.90 vs. 2.53%, respectively).

Conclusion:

The present study revealed that prepartum buffalo mammary gland secretion is accumulated pre-calving and stored inside the gland without any reflex to be milked. The response of mammary gland to prepartum milking varied between animals from moderate to no secretion. Prepartum buffalo mammary secretion can be identify as sticky secretion high in protein and low in fat and lactose. Precolostrum chemical composition dramatically differs from colostrum. Ovarian mammary gland axis seems to be a major determinant of the lactogenic activity of the buffalo mammary gland especially during its transformational epochs from precolostrum to colostrum. Further researches are needed to elucidate the buffalo ovarian dynamics related to the mammary gland function.

ARTICLE INFO

Article history:

Received 28 September 2015

Accepted 15 December 2015

Available online 24 December 2015

ACKNOWLEDGMENTS

The authors are grateful for the generous fund given by Prof. Dr. Mamdouh A. Sharafaldin, Professor of Animal Husbandry, Faculty of Agriculture, Cairo University and the Chairman of Board of Directors, Cairo Poultry Company (CPC), Egypt.

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(1) Nancy N. Kamel, (2,3) Yassein M. Hafez, (2) Ahmad F. El-Kholy, (1) Yahya A. Maareck and (1) Gamal A. Abou Ward

(1) Department of Animal Production, National Research Center, Postal Code 12262, Dokki, Giza, Egypt.

(2) Department of Animal Production, Faculty of Agriculture, Cairo University, Postal Code12613, Giza, Egypt.

(3) Hormones Lab., Cairo University Research Park. Postal Code12613, Giza, Egypt.

Corresponding Author: Nancy N. Kamel, Department of Animal Production, National Research Center, Postal Code 12262, Dokki, Giza, Egypt.

E-mail: nancy.nk3@gmail.com

Table 1: Mean yield and composition
of buffalo precolostrum and colostrum.

Type of               yield                Composition %
secretion          (kg/milking)
                                                Fat

Precolostum   0.24 [+ or -] 0.02 (b)   2.53 [+ or -] 0.10 (b)
Colostrum     3.37 [+ or -] 0.19 (a)   7.90 [+ or -] 0.31 (a)

Type of       Composition %
secretion
                      Protein                  Lactose

Precolostum   10.97 [+ or -] 0.12 (a)   2.61 [+ or -] 0.09 (b)
Colostrum     6.44 [+ or -] 0.22 (b)    4.18 [+ or -] 0.08 (a)

Type of       Composition %
secretion
                       TS *                      SNF *

Precolostum   17.40 [+ or -] 0.17 (b)   14.87 [+ or -] 0.12 (a)
Colostrum     19.97 [+ or -] 0.37 (a)   12.08 [+ or -] 0.23 (b)

Type of         Composition %              ER
secretion
                     Ash                units **

Precolostum   1.33 [+ or -] 0.06   349 [+ or -] 6 (b)
Colostrum     1.46 [+ or -] 0.04   444 [+ or -] 6 (a)

* TS: total solids and SNF: solids not fat.
** Electrical resistance measured in units.
Means within the same column having different
superscripts differ (P<0.0001).

Table 2: Yield and composition of buffalo
precolostrum relative to parturition.

Days           n            Yield              Composition %
relative to               (kg/day)
parturition                                         Fat

(-32 to -34)   3   0.21 [+ or -] 0.07 (b)    2.52 [+ or -] 0.45
(-29 to -31)   6   0.23 [+ or -] 0.06 (b)    2.01 [+ or -] 0.32
(-25 to -28)   6   0.13 [+ or -] 0.04 (b)    2.29 [+ or -] 0.29
(-21 to -24)   7   0.15 [+ or -] 0.04 (b)    2.45 [+ or -] 0.26
(-18 to -21)   8   0.17 [+ or -] 0.06 (b)    2.67 [+ or -] 0.27
(-14 to -17)   8   0.24 [+ or -] 0.06 (b)    3.10 [+ or -] 0.28
(-11 to -14)   8   0.23 [+ or -] 0.06 (b)    2.66 [+ or -] 0.29
(-7 to -10)    8   0.22 [+ or -] 0.05 (b)    2.48 [+ or -] 0.33
(-4 to -7)     9   0.30 [+ or -] 0.05 (ab)   2.72 [+ or -] 0.40
(-1 to -3)     9   0.47 [+ or -] 0.07 (a)    2.19 [+ or -] 0.25

Days                Composition %
relative to
parturition            Protein                    Lactose

(-32 to -34)   11.47 [+ or -] 0.40 (a)    1.96 [+ or -] 0.29 (c)
(-29 to -31)   10.90 [+ or -] 0.60 (ab)   2.07 [+ or -] 0.12 (bc)
(-25 to -28)   10.92 [+ or -] 0.34 (ab)   2.58 [+ or -] 0.39 (bc)
(-21 to -24)   11.16 [+ or -] 0.19 (ab)   2.52 [+ or -] 0.29 (bc)
(-18 to -21)   11.35 [+ or -] 0.28 (a)    2.16 [+ or -] 0.20 (bc)
(-14 to -17)   11.53 [+ or -] 0.39 (a)    2.29 [+ or -] 0.18 (bc)
(-11 to -14)   11.09 [+ or -] 0.43 (ab)   2.63 [+ or -] 0.18 (bc)
(-7 to -10)    10.84 [+ or -] 0.33 (ab)   2.69 [+ or -] 0.22 (bc)
(-4 to -7)     10.83 [+ or -] 0.34 (ab)   2.84 [+ or -] 0.24 (b)
(-1 to -3)     10.07 [+ or -] 0.28 (b)    3.63 [+ or -] 0.26 (a)

Days              Composition %
relative to
parturition           TS *                    SNF *

(-32 to -34)   16.87 [+ or -] 0.44   14.35 [+ or -] 0.39 (b)
(-29 to -31)   16.09 [+ or -] 0.57   14.08 [+ or -] 0.35 (b)
(-25 to -28)   17.21 [+ or -] 0.56   14.91 [+ or -] 0.45 (ab)
(-21 to -24)   17.45 [+ or -] 0.45   15.00 [+ or -] 0.34 (ab)
(-18 to -21)   17.05 [+ or -] 0.37   14.39 [+ or -] 0.25 (b)
(-14 to -17)   17.64 [+ or -] 0.29   14.54 [+ or -] 0.23 (ab)
(-11 to -14)   17.57 [+ or -] 0.43   14.92 [+ or -] 0.25 (ab)
(-7 to -10)    17.50 [+ or -] 0.59   15.01 [+ or -] 0.37 (ab)
(-4 to -7)     17.91 [+ or -] 0.63   15.20 [+ or -] 0.38 (ab)
(-1 to -3)     17.86 [+ or -] 0.56   15.68 [+ or -] 0.41 (a)

Days                Composition %            ER units **
relative to
parturition              Ash

(-32 to -34)   0.92 [+ or -] 0.29 (b)    317 [+ or -] 29 (b)
(-29 to -31)   1.11 [+ or -] 0.21 (b)    325 [+ or -] 18 (b)
(-25 to -28)   1.42 [+ or -] 0.21 (b)    346 [+ or -] 25 (ab)
(-21 to -24)   1.32 [+ or -] 0.14 (b)    354 [+ or -] 17 (ab)
(-18 to -21)   0.99 [+ or -] 0.18 (b)    344 [+ or -] 20 (ab)
(-14 to -17)   0.96 [+ or -] 0.22 (b)    348 [+ or -] 15 (ab)
(-11 to -14)   1.19 [+ or -] 0.24 (b)    340 [+ or -] 8 (ab)
(-7 to -10)    1.48 [+ or -] 0.13 (ab)   345 [+ or -] 19 (ab)
(-4 to -7)     1.52 [+ or -] 0.11 (ab)   348 [+ or -] 16 (ab)
(-1 to -3)     1.97 [+ or -] 0.09 (a)    393 [+ or -] 24 (a)

* TS: total solids and SNF: solids not fat.
** Electrical resistance measured in units.
Means within the same column having different
superscripts differ (P<0.05).

Table 3: Yield and composition of buffalo
colostrum relative to parturition.

Hours         n        Yield (kg/day)            Composition %
relative to
parturition                                           Fat

0             10   5.18 [+ or -] 0.63 (a)   6.97 [+ or -] 0.92 (b)
12            10   2.51 [+ or -] 0.44 (b)   10.28 [+ or -] 0.92 (a)
24            10   2.63 [+ or -] 0.61 (b)   8.74 [+ or -] 0.83 (ab)
36            10   2.40 [+ or -] 0.55 (b)   8.03 [+ or -] 0.74 (ab)
48            10   3.08 [+ or -] 0.50 (b)   8.20 [+ or -] 0.71 (ab)
60            10   3.46 [+ or -] 0.30 (b)   7.49 [+ or -] 1.03 (b)
72            10   3.82 [+ or -] 0.46 (b)   6.37 [+ or -] 0.78 (b)
84            10   3.44 [+ or -] 0.26 (b)   7.17 [+ or -] 0.72 (b)

Hours         Composition %
relative to
parturition           Protein                    Lactose

0             10.14 [+ or -] 0.35 (a)    3.81 [+ or -] 0.12 (cd)
12            8.10 [+ or -] 0.45 (b)     3.57 [+ or -] 0.15 (d)
24            6.30 [+ or -] 0.34 (c)     3.88 [+ or -] 0.22 (cd)
36            5.34 [+ or -] 0.25 (cd)   4.06 [+ or -] 0.29 (bcd)
48            5.56 [+ or -] 0.30 (cd)   4.23 [+ or -] 0.21 (abcd)
60            5.59 [+ or -] 0.40 (cd)    4.65 [+ or -] 0.27 (ab)
72            5.43 [+ or -] 0.27 (cd)   4.49 [+ or -] 0.27 (abc)
84            5.00 [+ or -] 0.21 (d)     4.74 [+ or -] 0.19 (a)

Hours         Composition %
relative to
parturition            TS *                      SNF *

0             22.68 [+ or -] 1.07 (a)   15.71 [+ or -] 0.36 (a)
12            23.55 [+ or -] 1.01 (a)   13.28 [+ or -] 0.49 (b)
24            20.22 [+ or -] 1.04 (b)   11.49 [+ or -] 0.58 (c)
36            18.67 [+ or -] 0.61 (b)   10.64 [+ or -] 0.21 (c)
48            19.26 [+ or -] 0.63 (b)   11.06 [+ or -] 0.49 (c)
60            19.29 [+ or -] 1.09 (b)   11.77 [+ or -] 0.63 (c)
72            17.79 [+ or -] 0.77 (b)   11.42 [+ or -] 0.33 (c)
84            18.35 [+ or -] 0.56 (b)   11.18 [+ or -] 0.30 (c)

Hours              Composition %           ER units **
relative to
parturition             Ash

0              1.76 [+ or -] 0.07 (a)    438 [+ or -] 19
12            1.61 [+ or -] 0.10 (ab)    437 [+ or -] 18
24            1.31 [+ or -] 0.14 (bc)    433 [+ or -] 23
36             1.24 [+ or -] 0.08 (c)    442 [+ or -] 24
48            1.27 [+ or -] 0.14 (bc)    435 [+ or -] 12
60            1.53 [+ or -] 0.14 (abc)   453 [+ or -] 12
72            1.51 [+ or -] 0.14 (abc)   438 [+ or -] 19
84            1.44 [+ or -] 0.08 (abc)   479 [+ or -] 16

* TS: total solids and SNF: solids not fat.
** Electrical resistance measured in units.
Means within the same column having different
superscripts differ (P<0.05).
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Author:Kamel, Nancy N.; Hafez, Yassein M.; Kholy, Ahmad F. El-; Maareck, Yahya A.; Ward, Gamal A. Abou
Publication:Advances in Environmental Biology
Article Type:Report
Date:Dec 1, 2015
Words:5482
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