Lactation Curves for First Lactation Egyptian Buffalo
G. H. METRY and K. A. MOURAD
ABSTRACT
First lactation records of 1538 buffalo
maintained at the Animal Production Re-
search Institute fanns in 1967 to 1990
were used to determine lactation curves
for three lactation lengths: >28 and <150
d, >149 d, and ~308 d, as well as all
records. Daily milk yields were summed
by 14-d intervals for analyses. Yields
peaked at the first, fifth, sixth, and sev-
enth periods for >28 and <150 d, >149 d.
~308 d, and all records. Herd-year-
season significantly affected milk yield
in all periods. Persistency for all records
and three groupings (>28 d, >149 d, and
~308 d in milk) was highest for the
~308-d group (1.02 vs..85 for >149 d,
.57 for >28 d, and .47 for all records).
Herd-year-season of calving significantly
affected persistency in all records and
the three subset groupings. Persistency
was higher for buffalo calving in spring
and summer for all records, records >28
d, and records> 149 d but in summer and
autumn for the ~308 d records. In
records ~308 d, the correlation coeffi-
cient between persistency and milk yield
(r = .06) was not significant but was
negative with season of calving (r =
-.11).
(Key words: buffalo, first lactation,
curve)
INTRODUCTION
Buffalo (Type Beheri of genera Bubalis) are
considered to be the principal milk-yielding
Received July 2, 1993.
Accepted December 22. 1993.
1994 J Dairy Sci 77:1306-1314
Animal Production Research Institute
Agriculture Research Center
Ministry of Agriculture
Giza, Egypt
J. C. WILK and B. T. McDANIEL
Department of Animal Science
North Carolina State University
Raleigh 27695-7621
species in Egypt and provide milk and milk
products that are important sources of protein,
energy, vitamins, and minerals for the diet of
the Egyptian people. However, the amount of
milk yield in Egypt is inadequate to meet the
needs of the people there. A better understand-
ing of factors influencing yield of buffalo is
needed to increase the total amount of milk
produced in Egypt. Animal health, feeding,
and reproductive programs and their manage-
ment account for 75% of variation in herd
yield, and the remaining 25% is due to genetic
sectors (3). The rate of secretion of milk in
dairy animals displays a definite trend through-
out the lactation period. Following parturition,
daily milk yield increases to a peak and then
gradually declines until the lactation is ended.
Some lactations are terminated or truncated
before the desired time (usually 305 d) because
of low persistency (6). The shape of the lacta-
tion curve is influenced by parity and season
of calving in cattle (25). Fann, parity, and
season of calving influenced the shape of lacta-
tion curve of Egyptian buffalo (16).
Qualitative and quantitative differences in
feeding and management are among the impor-
tant causes of variation in milk yield (18) and
greatly influence the shape of the lactation
curve. The shapes of lactation curves could be
measured by the persistency of yield [i.e., the
ratio of yields in consecutive periods (4)].
Maymone and Malossini (11) estimated the
persistency of milk yield in buffalo as the ratio
of amount of milk produced in the second 100
d of lactation to that produced in the first 100
d of lactation. Most studies of milk yield of
buffalo have used only lactations of ~305 d (1,
17), but some have used lactations >150 d (16).
The objectives of this study were to inves-
tigate the effects of herd-year-season and age
at calving on the shape of the production curve
1306
 LACTATION CURVES 1307
of first lactation buffalo and their persistency persistency of lactations of variable length >28
in lactations of different lengths. d, intennediate length> 149 d, and those ~308
d. Lactations >28 d included those >149 d, and
MATERIALS AND METHODS those >149 d included those ~308 d.
All records were used to detennine the
Daily milk yield records were available pooled lactation curve with the following
from 1538 first lactation buffalo calving from model using software of SAS (19).
1967 to 1990 in herds at three experimental
stations of the Animal Production Research
Institute, (El-Nattaf EI-Gedid, EI-Nattaf EI- Yijklm = (30 + HYSi + bl(Pj) + b2(P~)
Kadem, and Mehallet Mousa, Egypt). The
+ b3(P~) + ~jklm [1]
herds were maintained according to feeding
and management standards adopted by the in-
stitute. During winter and spring (December to where
May), cows were grazed on Egyptian clover as
the only source of feed. During summer and Yijklm = 14-d milk yield record,
autumn (June to November), cows were kept {30 = intercept,
under open sheds covered with asbestos roofs HYSi = random effect of herd-year-
4 m high. During this period, feeding consisted season i,
of a concentrate mixture, wheat or rice straw, bl = partial linear regression of milk
and a limited amount of clover hay when yield on stage of lactation (peri-
available. The cows were milked by hand od),
twice daily at 0730 and 1630 h without the ~ = partial quadratic regression of
presence of the calf. In order to include all milk yield on (stage)2,
possible variations in the shape of lactation b3 = partial cubic re¥ression of milk
curve, records were included whether or not yield on (stage) ,
308 d of lactation were completed. Pj = stage of lactation j (j = I, 2,
Preparatory to this study, the daily milk 3... ,22),
yields were summed by l4-d intervals for the Pk = (stage)2,
duration of the lactation. A total of 498 of the PI = (stage)3, and
1538 records had <29 d of lactation and were eijklm = a random residual, normally and
deleted from some analyses because of prob- independently distributed with
lems occurring before, at, or just after parturi- zero mean and vanance· 2
(Je'
tion. The remaining records were then sorted
into four groupings: 1040 lactations >28 d, 344
The records of the three groups based on
lactations >28 and <150 d, 651 lactations >149
lactation length (>28 d but <150 d, >149 d, and
d, and 260 lactations ~308 d (22 periods) (Ta-
~308 d) were used to detennine the lactation
ble I). The subsets were for estimation of
curves. The three subsets were analyzed as
follows.

TABLE I. Mean milk yield and lactation length in the = {30+ HYSi + b 1(Gj x Pk)
three different groupings and all records. + b2(Gj x ph
+ b3(Gj x P~)
Mean + eijklmn
Record milk Lactation [2]
length yield length
(d) (n) (kg) (d) where
>28 1040 892 212
>28 and <:150 1
>149
344
651
360
1229
88
290
Yijklmn = 14-d milk yield record,
~308 260 1661 393 (Gj x Pk) = stage of lactation k (k =
Total 1538 606 144 1,2,...22) within group j (j =
1,2,3),
lThe group included to draw a separate curve.

Journal of Dairy Science Vol. 77, No.5, 1994

1308 METRY ET AL.

(OJ x P~) = (stageY within group, A2

k
= (ageY.
(OJ x P~) = (stage)3 within group, A3
I
= (age>3,
and other descriptions are as in Model [I]. and other descriptions are as in Model [I].
To determine the lactation curves by season The effect of age at calving on milk yield
of calving from all records, the following by stage of lactation by season was calculated
model was used. from the following model.

Yijklmn = 130+ HYj + bl(Sj x Pk) Yijldmn = f30 + HYi + Sj + bl(~) + b2(Ah
+ b2(Sj x ph
+ b3(Sj x P~) + b3(A~) + b4(Sj x Ak)
+ eijklmn [3]
+ bs(Sj x Ar) + b6(S x A3)
where + eijldmn [5]

where
Yjjklmn = 14-d milk yield record,
HYj = random effect of herd-year i, 14-d milk record,
bl = partial linear regression of season j of calving,
milk yield on stage within
season, partial linear regression of
b2 = partial quadratic regression milk yield by stage on age at
of milk yield on (stageY calving.
within season, b2 = partial quadratic regression
b3 = partial cubic regression of of milk yield by stage on
milk yield on (stage)3 within (ageY,
season. b3 = partial cubic regression of
(Sj x Pk) = stage of lactation k within milk yield by stage on (age)3,
season j of calving, [1 = b4 = partial linear regression of
spring (March to May), 2 = milk yield by stage on age at
summer (June to August), 3 = calving by season,
autumn (September to bs = partial quadratic regression
November), and 4 = winter of milk yield by stage on
(December to February)], (ageY by season,
(Sj :-: ph = (stageY within season, b6 = partial cubic regression of
milk yield on (age)3 by sea-
(Sj x P 3)
m
= (stage)3 within season, and son,
Ak = age k at calving in mo,
with f30 and eijldmn as described earlier. Ar = (ageY,
The effect of age at calving on milk yield 3
Am = (age)3,
for each stage of lactation (14 d milk yield)
(Sj x Ak) = age k at calving by season j,
was obtained from the following model.
(Sj x Ah = (ageY by season.
Yijklm = 130 + HYS j + bl(Aj) + ~(A~) (Sj x A~) = (age>3 by season,

+ b3(A?) + ~jklm [4]

and other descriptions are as in Model [4].
Persistencies in all records and the three
where classes (>28, >149, and ~308 d of lactation)
were each estimated as the ratio of milk yield
Yijklm = 14-d milk record, in the second 100 d of lactation to milk yield
Aj = age jat calving in months in the fIrst 100 d of lactation (P2:1) (11) using
G = 27,28,.... ,77), Model [6],

Journal of Dairy Science Vol. 77, No. S. 1994

 LACTAnON CURVES 1309

Yijklmn = t30 + HYi + Sj + bl(~) TABLE 2. Regression of milk yield! by 14-d periods on
age at calving, (age)2, (age)3, and percentage of variance
+ b2(Ar> + b3(A~) + b4(Sj x Ak) due to age at calving within herd-year-season (HYS) for
+ bs(Sj x At> + b6(S x A3)
first lactation buffalo.

Variance
+ eijklmn [6] due to
Periods 2 age within
where in lactation Age (Age)2 (Age)3 HYS
(%)
Yijklm = the persistency percentage, I 12.9** -.28** .002** .79
2 14.7** -.33** .002** .38
and other descriptions are as in Models [4] and 3 -9.7 .22 -.002 .32
[5]. 4 -14.1 .36 -.003 .53
S -3.9 .11 -.001 .64
6 -8.8 .23 -.002 .66
RESULTS AND DISCUSSION 7 -10.7 .25 -.002 .41
8 -197 .46 -.003 .76
The mean milk yield, lactation length, and 9 -10.2 .26 -.002 .24
numbers of records for the subsets and overall 10 -7.4 .18 -.001 .56
II -10.9 .27 -.002 .75
records are given in Table l. Of the total 12 -14.7 .36 -.003 .48
records, 32% were <29 d and were excluded 13 -13.8 .33 -.003 .90
from analyses because of problems before, at, 14 -14.9 .36 -.003 .61
and after parturition (abortion, stillbirth, or IS -16.9 .40 -.003 .75
failed to lactate). Only 16.9% of the 1538 fIrst 16 -6.2 .13 -.001 .52
17 -18.5 .44 -.003 .82
lactation buffalo completed lactations that 18 1.0 -.04 .001 1.13
were ~308 d in length. The mean milk yield 19 -3.9 .08 -.001 1.03
was nearly doubled, from 892 kg for the 1040 20 5.3 -.12 .001 1.96
records >28 d to 1661 kg for the 260 records 21 .8 -.03 .000 1.94
~308 d of lactation. Also, the mean lactation
22 6.8 -.15 .001 1.27
length increased from 212 to 393 d for the **p < .01.
respective groups of records. lAdjusted for HYS.
214-d Milk yield.
Effect of Age at Calving

Regressions of milk yield (adjusted for

herd-year-season) from Model [4] by 14-d
periods on age, (ageY, and (ageY at calving
and percentage of variance because of age at
calving within herd-year-season from Model
[4] are in Table 2. The effects of age at
calving, (ageY, and (age)3 were signifIcant (P <
.01) for the first and second periods but not for
periods 3 through 22. This result agrees some-
what with results of Singh and Yadav (20),
who reported that mean milk yield by Murrah
buffalo in India was not affected by age at fIrst
calving. The percentage of variance because of
age at calving within herd-year-season was
small «1%) for early stages but increased to
1.27 by period 22 (Table 2).
The effects of herd-year, season, and age at
calving from Model [5J are shown in Table 3.
Herd-year had a signifIcant effect (P < .01) on
periods 1 through 16 (Table 3) but was not
significant from periods 17 to 22. The herd-
year effect reflected the fluctuation in feed
supply between years that was due to climatic
conditions. These results are similar to those
obtained in other studies of buffalo (16,20,24)
and cattle (9, 22), but disagreed with observa-
tions by Khosla et al. (7), who reported that
year of calving had no signifIcant effect on
daily milk yield of first lactation buffalo in
Pakistan. Their fIndings might have been in-
fluenced by the few years (1978 to 1980) co-
vered by their study.
Season of calving (Model [5]) was not sig-
nificant for the 22 periods in milk. However,
age at calving, (ageY, and (age)3 were signifi-
cant (P < .01) in periods 1, 2, and 4 but not
significant in periods 3 and 5 through 22 (Ta-
ble 3). The percentage of variance because of
periods within season was very small, .004,
but was .11 for periods within herd-year. The

Journal of Dairy Science Vol. 77, No.5, 1994

1310 METRY ET AL.

TABLE 3. Analysis of variance for factors affecting stage of lactation (14 d of lactation).

Lactation periods l
Source df 2 3 4 5 to 16 17 to 22
F-value
Herd-year 49 4.2** 5.8** 3.3** 3.1** 1.3 - 2.0** NS
Season 3 1.1 1.1 1.4 1.3 NS NS
Age at calving 1 8.8** 3.3 t 1.9 2.7 NS NS
(Age)2 I 7.9** 2.91 1.5 3,P NS NS
(Age)3 1 7.0** 2.4 1.2 3.6 t NS NS
(Age) x season 3 1.1 1.1 1.3 1.3 NS NS
(Age)2 x season 3 1.2 1.1 1.2 1.3 NS NS
(Age)3 x season 3 1.2 1.0 1.1 1.3 NS NS
R2, % 17 18 16 16 14 - 18 IS - 23

1I4-d Milk yield,

tp < .10.
*p < .05.
"p < .01.

effects of age at calving, (ageY, and (ageY lactation Curves

within season were not significant for all peri-
ods. Periods in milk were affected by factors
other than season and age at calving, such as
feeding, managerial practices, or the weather.
Cockrill (4) reported that the effect of tempera-
ture and humidity were of great importance for
milk yield in buffalo. Buffalo are more sensi-
tive than cattle to heat exposure. As environ-
mental temperatures rose, water consumption
increased, but, if the feed intake was
depressed, milk secretion decreased. Misra et
al. (14) found that both food and water intakes
in buffalo were reduced by ambient tempera-
tures >32°C. Our findings agreed with those
for Sahiwal cows of India (2) where season of
calving had no significant effect on the weekly
milk yield in the last few weeks of lactation.
For Egyptian buffalo, Alim (1) and Salem (17)
found that 305-d milk yield in multiple lacta-
The estimates obtained from Model [1]
were used to locate the points of the pooled
curve (Figure 1). Estimates from Model [2]
were used to obtain overall curves of expected
yield for periods 1 through 12 for records >28
but <150 d and 22 for records >149 d and
~308 d (Figure 2). The pooled curve peaked at
periods 7 and 8, decreased steadily until period
19, and then increased to period 22. The con-
figuration of the lactation curve of records >28
but <150 d reached its peak at period I, main-
l00r--------------~
80

tions of buffalo was not affected by season of ,...

calving. However, the results were not similar
for Egyptian buffalo (5, 14) and for buffalo in
Pakistan (7).
Proportions of variance within the 22 peri-
ods attributable to the fifth analytical model
ranged from R2 = .11 to .30 (fable 3). The 20f-
contribution of herd-year to total variance
within periods was 10 to 17%. These are less
than the 30 to 38% of the total variance in 4 6 8 10 12 14 16 18 20 22

milk yield by cattle in Brazil (12). The percent- p.,locta

age of variance attributable to season of calv- Figure I, Overall lactation curve over 22 14-<1 periods
ing was relatively small at .54%. for first lactation buffalo.

Journal of Dairy Science Vol. 77. No. S. 1994

 LACTATION CURVES
100,-----------------,
20
\
2 4 6 8 10 12 14 16 18 20
periods
Figure 2. Lactation curves over 22 14-d periods for
flrstlactation buffalo with >28 d (0), >149 d (+), and ~308
d (.) of lactation.
tained this peak for three additional periods,
and decreased rapidly until period 12, which
reflected the low persistency. Records >149 d
and ~308 d peaked at periods 5 and 6, respec-
tively, and remained near the same level
through periods 6 and 8 and then decreased
until periods 21 and 22. The rate of decline in
milk yield from the peak period to the next one
was 5.0, 1.4, 1.3, and 1.5% for records >28 but
<150 d, >149 d, >308 d of lactation, and all
records, respectively. The mean rate of decline
from the peak to period 12 was 22.1 % for
records >28 but <150 d, but was only 3.1,2.8,
and 1.3%, respectively, from peak to period 22
for records >149 d, ~308 d, and all records.
Rates of decrease were less than reported for
crossbred cattle in India (15). The rapid rate of
decline in records >28 but <150 d and records
>149 d was probably due to the short lactation
records included, which terminated before 308
d of lactation. This result agrees with that of
Singh et a1. (22), who reported that short lacta-
tions «305 d) were more common than long
lactations (>305 d) for all parities of Murrah
buffalo (956 short lactations out of 1609
records).
Milk yield curves (Model [3]) for lactations
initiated by calvings in the same season within
herd-year are shown in Figure 3. The shape of
the yield curve following winter calvings (De-
cember to February) was quite different from
those of other seasons. The peak was similar to
that for the other calving seasons. The ascend-
1311
100'1- - - - - - - - - - - -
i
80
::
-g 60
'iia.
'6
:!.. 40
:!!
E
20 ~
I
oLI- _
22 2 4 6 8 10 12 14 16 18 20 22
periods
Figure 3. Lactation curves over 22 14-d periods for
seasons of calving spring e-).
summer (+). autumn (.), and
winter (0) for first lactation buffalo.
ing phase was seven periods; then yield
declined rapidly from periods 8 to 19 and
increased again from periods 20 to 22. Lacta-
tion rates following spring, summer, and au-
tumn calvings remained at near peak levels for
2, 3, and 2 periods and declined steadily there-
after until periods 19, 21, and 22, respectively.
The yield trend for winter calving was in-
fluenced between periods 8 and 19 by changes
from winter feeding regimens (surplus energy
and protein for milk yield from green clover,
and mild weather) to summer feeding regimens
(dry feed with high fiber and deficient protein,
heat stress, and abrupt change from green to
dry feed) and increased again at the beginning
of the next winter season. This result agrees
with reports that the differences in lactation
curves of Flekvieh cattle in Hungary between
calving seasons reflected differences in feeding
(8). The trend following winter calvings agreed
with that for imported Friesian cattle in Nige-
ria (23). Also, Madalena et al. (10) reported
significant season effects on the parameters of
the lactation curves of Holsteins in Brazil.
Cows that calved in the wet season in Brazil
had higher initial yield but were less persistent
than those cows calving in the dry season.
Persistency
Persistencies (Model [6]) of milk yield as
the ratio of the second 100 to the first 100 d of
milk yield are presented in Table 4. The
records with ~308 d of lactation had 102%
Journal of Dairy Science Vol. 77. No. S. 1994
1312 METRY ET AL.

TABLE 4. Least squares means and standard errors for persistency! of first lactation buffalo calving in different seasons.

Lactation records
Items >28 d >149 d <?:308 d All records
X SE X SE X SE X SE
Overall .57 .13 .85 .11 1.02 .10 .46 .10
Spring .62 .03 .88 .03 1.01 .03 .52 .03
Summer .64 .04 .89 .03 1.11 .04 .54 .03
Autumn .54 .03 .87 .03 1.06 .03 .41 .03
Winter .48 .03 .77 .02 .90 .03 .38 .03

1A ratio of the second 100 and first 100 d of lactation.

persistency compared with persistencies of in records ~308 d of lactation were 10 I, Ill,

85% in lactations >149 d, 57% in lactations
>28 d, and only 46% in all records. A similarly
calculated persistency of 83% was reported for
first lactation buffalo in Italy (4). Records ~308
d of lactation had information in all periods,
but records >28 d, >149 d, and all records had
substantial numbers of records terminated
earlier than 308 d of lactation. In addition, a
higher proportion of the records in the >28 d,
>149 d, and all records lactation groupings
may have been influenced by poor design and
management of health, nutrition, feeding, and
reproduction programs. Also, lactations were
terminated early because buffalo in poor condi-
tion at calving were unable to maintain accept-
able milk yield.
Mean persistencies for cows calving in
spring, summer, autumn, and winter calvings
106, and 90%, respectively. Trends were simi-
lar for lactations >28 d, >149 d, and all records
even though persistency was much lower. The
similarity of trends indicates that the seasonal-
ity of production may be stimulated by the net
effect of the season of year in which parturi-
tion occurred (16). The season stimulus is a
consequence of nutrition, health, and climate
and shows its influence at all stages of the
lactation.
Herd-year had a highly significant effect (P
< .01) on persistency for all records, those >28
d, and those >149 d, and this effect was sig-
nificant (P < .05) for the ~308-d grouping
(Table 5). Season of calving affected (P < .05)
the persistency of all records and of records
>28 d (P < .01). Season was significant (P <
.10) for records ~308 d but not for records

TABLE 5. Analysis of variance for persistencyl of lactation milk yield for records >28 d, >149 d, <?:308 d of lactation,
and all records.

Lactation records

28 d >149 d <?:308 d All records

Source df MS df MS df MS df MS
Herd-year 49 1.15 u 47 .18-- 28 .05- 49 2.17--
Season 3 1.19-- 3 .12 3 .o7 t 3 .39*
Calving age 1 .30 1 .16-
(Age)2 1 .30 1 .17-
(Age)3 1 .27
Age x season 4 .12 3 .06 4 .30 t
(Age)2 x season 4 .12 3 .06
(Age)3 x season 4 .18*
Residual 984 .16 588 .07 220 .03 1481 .14
R2, % 28.9 222 30.4 35.7

!A ratio of the second 100 and first 100 d of lactation.

tp < .10.
-p < .05.
up < .01.

Journal of Dairy Science Vol. 77. No.5, 1994

 LACTATION CURVES
>149 d of lactation. These findings were simi-
lar to those of Zamorano Villarreal (26) with
Friesian cows in Mexico. Age at calving had
significant linear and quadratic effects (P <
.05) on the persistency in ~308-d records but
was not significant in >28-d records (Table 5),
which agrees with findings (26) for cattle. This
discrepancy could be explained by the wide
variation in age at calving (27 to 77 mo) in the
>28-d grouping; consequently, persistency var-
ied.
Correlation was positive but not significant
between persistency and 308-d milk yield (r =
.06), but negative between persistency and sea-
son of calving (r = -.11). However, the magni-
tude of correlation between persistency and
milk yield was less than for first lactation
Holstein-Friesians (r =
.35), which disagrees
with Zamorano Villarreal (26).
CONCLUSIONS
Factors affecting the shape of lactation
curves of buffalo are likely similar to those of
cattle. A high percentage of the first lactation
buffalo in this study had short lactations
(37.4% had <150 d of lactation). Short lacta-
tions and the consequent low yearly milk
yields drastically reduce the usefulness of
buffalo as dairy animals. More research to
identify the underlying genetic and environ-
mental causes should be a high priority for
areas where dairy buffalo are used. Seasonal
differences suggest that improved management
through adequate feeding for milk yield, dis-
ease prevention, parasite control, protection
against extremes of climate, and a regular
reproductive timetable could be useful to im-
prove milk yield.
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