Near-Infrared Spectroscopy for Dairy Management: Measurement
of Unhomogenized Milk Composition
*Department of Environmental Information and Bio-Production Engineering,
ABSTRACT
The potential of near-infrared spectroscopy to mea-
sure fat, total protein, and lactose contents of unhomog-
enized milk was studied for use in dairy management,
as a new tool for on-line milk analysis in the process
of milking. Influence of the spectral region, sample
thickness, and spectral data treatment on the accuracy
of determination was investigated.
Transmittance spectra of 258 milk samples, collected
at different stages of the milking process, were obtained
with a spectrophotometer (NIRSystems 6500; FOSS-
NIRSystems, Silver Spring, MD) in the wavelength
range from 400 to 2500 nm with sample thicknesses of
1 mm, 4 mm, and 10 mm.
The spectral region and sample thickness were found
to be significant factors for milk fat and total protein
determination but not the lactose determination. The
best accuracy was obtained with the 1100 to 2400 nm
region, 1-mm sample thickness, and the first derivative
data transformation. For the spectral region from 700
to 1100 nm, close accuracy was obtained for fat with a
10-mm sample and for total protein with a 1-mm sample
thickness. The sample thickness did not change signifi-
cantly the accuracy of lactose determination. Different
treatments of spectral data did not improve the calibra-
tions for fat and protein.
For the region from 700 to 1100 nm, where inexpen-
sive on-line sensors could be used, the highest positive
coefficients for fat were at 930, 968, 990, 1026, 1076,
and 1092 nm; for lactose were at 734, 750, 786, 812,
908, 974, 982, and 1064 nm; and for total protein were
at 776, 880, 902, 952, and 1034 nm.
(Key words: unhomogenized milk composition, near-
infrared spectroscopy)
Received October 15, 1998.
Accepted June 21, 1999.
1999 J Dairy Sci 82:2344–2351 2344
R. TSENKOVA,* S. ATANASSOVA,† K. TOYODA,*
Y. OZAKI,‡ K. ITOH,§ and T. FEARN储
Kobe University, Kobe 657, Japan
†Department of Mathematics and Physics,
Thracian University, Stara Zagora 6000, Bulgaria
‡Department of Chemistry,
Kwansei Gakuin University, Nishinomiya 662, Japan
§Laboratory of Agricultural Process Engineering,
Hokkaido University, Sapporo 060, Japan
储Department of Statistical Science,
University College London, London WC1E 6BT, United Kingdom
Abbreviation key: CV = coefficient of variation, CVr =
cross-validation correlation coefficient, NIR = near-in-
frared, PLS = partial least square, SECV = standard
error of cross-validation, T = transmittance.
INTRODUCTION
Milk composition is an important factor in dairy
farming. Its measurement is essential for the efficient
utilization of cows, cattle breeding, and dairy industry.
Fat and total protein content of milk have economic
importance because, in most countries, milk trade is
based on these components and the somatic cell count.
Daily measurement of milk composition is becoming a
very important information source for dairy manage-
ment decisions and could be a valuable diagnostic tool
as well (15). Decrease in the milk fat content could be
an indicator for imbalance in the forage to concentrates
ratio in the diet (6). Lactose content of milk is usually
very stable. Its deviation could be used as an additional
indicator for mastitis diagnosis. Total protein content
of milk is an individual parameter for each cow, which
changes with and within the lactation stage and when
somatic cells count increases. Protein content of milk
could also be used as an energy supply indicator for
each cow. Low protein content of milk indicates energy
deficiency. With dairy management requirements for
high milk quality and the rapid development of milking
robots a clear necessity exists for a nondestructive,
rapid method and instrumentation for on-line milk
analysis. Near-infrared (NIR) technology is quite suit-
able for that purpose and especially the short wave-
length (near NIR) range because of the inexpensive
optical sensors available on the market.
Routine analytical methods used for milk composi-
tion measurement are destructive, expensive, time and
labor consuming, and off-line by nature. A widely used
instrument for milk analysis is Milko Scan (Foss-Elec-
 MILK COMPOSITION MEASUREMENT
tric A/S, Hillerod, Denmark), which is based on infrared
spectroscopy (1). Even the largest laboratories equipped
with a Milko Scan instrument can provide milk compo-
sitional data for samples taken from each cow once or
twice per month. The information obtained is expensive
and does not help every day management decisions at
dairy farm. Modern farm management involves auto-
mated systems to control the production process and
collection of on-line data such as milk yield and body
weight of the cows. The use of NIR spectroscopy for on-
line milk control on the farm could be a further step in
that concept (20). Near-infrared spectroscopy has been
demonstrated as a successful tool in agriculture and in
the food industry for rapid and accurate compositional
analysis and for quality control of different products
(7, 21). Its advantages include speed, simplicity, and
simultaneous nondestructive measurements of a num-
ber of constituents. The NIR spectroscopy has been used
to measure the content of various constituents in dairy
products such as milk, milk powder, whey, and cheese
(2, 3, 4, 5, 8, 9, 10, 11, 12, 22). Dry milk (2) and homoge-
nized milk samples (4, 12) have commonly been used
for milk analysis, which is inconvenient for on-line mea-
surements. Only a few authors (14, 16, 17, 18, 19, 20)
report analysis of unhomogenized milk. We studied the
potential of NIR spectroscopy to measure fat, total pro-
tein, and lactose contents of unhomogenized milk to be
used in the process of milking for dairy management
purposes. Influence of the spectral region, sample thick-
ness, and spectral data treatment (smoothing and de-
rivative transformation) on the accuracy of determina-
tion were investigated.
The aim of this study was to investigate the potential
of NIR spectroscopy for measurement of fat, total pro-
tein, and lactose in unhomogenized milk. If accurate,
this method could then be used for dairy management
as a new tool for on-line milk analysis in the process
of each milking. The accuracy of NIR spectroscopy de-
termination of milk composition in the near NIR region,
suitable for on-line measurements with fiber optic
probes and relatively inexpensive silicon detectors, was
compared with that of the 1100 to 2400 nm region,
which allows composition measurement with high accu-
racy. Investigation of the influence of sample thickness
and spectral data treatment on the accuracy of milk
composition determination for both spectral regions
was also carried out.
MATERIALS AND METHODS
Milk Samples
A total of 258 individual milk samples was collected
from three Holstein cows twice a month for a 6-mo
period. Equal portions of milk (3 L) were taken during
2345
the morning and evening milkings with a milk yield
meter (True Test; Alfa Laval Agri, Tumba, Sweden).
The experiment was carried out between the second
and the eighth month of lactation for cows that were
in their second lactation. All cows were kept on the
same diet during the experiment.
All samples were analyzed simultaneously with the
reference method (1) for fat, total protein, and lactose
contents by Milko Scan (Foss-Electric A/S, Hillerød,
Denmark).
NIR Spectra
Transmittance (T) milk spectra with path lengths of
1 mm, 4 mm, and 10 mm were obtained from all samples
with a spectrophotometer (NIRSystems 6500; FOSS-
NIRSystems, Silver Spring, MD) by using quartz cu-
vettes with 1 mm thick walls. Spectra were taken in
the wavelength range from 400 to 2500 nm at 2-nm
intervals and were recorded in the linked computer as
absorbance [i.e., log (1/T)]. Prior to spectral analysis,
each sample was warmed to 40°C in a water bath.
Spectral data were analyzed to obtain calibration
models in two regions: the region from 700 to 1100 nm
with 1-, 4-, and 10-mm path lengths and the region
from 1100 to 2400 nm with a 1-mm path length. In the
1100 to 2500 nm region, the absorptions for 4- and 10-
mm path lengths were very high, and the spectra were
noisy; hence, no calibrations were performed. The spec-
tral data for 1-mm sample thickness from 2400 to 2500
nm were also noisy and were not used.
Treatment of NIR Data
A commercial software program (Pirouette Version
2.0; Infometrics, Inc., Woodinville, WA) was used to
process the data and to develop models for determina-
tion of fat, total protein, and lactose contents. Calibra-
tion models for each combination of spectral region,
sample thickness, and data treatment were obtained.
The methods for data treatment included smoothing
the spectral data over different intervals, from 5- to 25-
point windows (10-50 nm), and first or second derivative
transformation of log (1/T) data with different window
sizes, from 5- to 25-point intervals. The smoothing and
derivative transformations were based on the Savitzki-
Golay second-order polynomial filter (13).
The calibration was performed using partial least
square (PLS) regression. The PLS utilized both the
spectra and the respective reference data for the exam-
ined samples to determine the latent variables (PLS
factors). In the development of all calibration models,
15 PLS factors were set up as maximum. The optimum
number of PLS factors used in the models was deter-
Journal of Dairy Science Vol. 82, No. 11, 1999
2346
TABLE 1. Range, mean, and standard deviation (SD) of milk compo-
nents in examined samples.
Parameter Minimum Maximum Mean
Fat, % 0.34 10.41 3.505
Protein, % 2.60 3.99 3.132
Lactose, % 3.94 4.74 4.399
mined by a cross-validation method. In cross-valida-
tion, five samples were temporarily removed from the
calibration set to be used for validation. With the rest
of the samples, a PLS model was developed and applied
to predict the respective milk component content of the
group of five samples. The results were compared with
the respective reference values. This procedure was re-
peated several times until a prediction for all samples
was obtained. Performance statistics were accumulated
for each group of removed samples. The validation er-
rors were combined into a standard error of cross vali-
dation (SECV), which was accepted as a measure of
the accuracy of determination. The optimum number
of PLS factors in each model was defined to be the one
that corresponded to the lowest SECV.
RESULTS AND DISCUSSION
The experiment was designed to follow the changes in
milk composition during lactation and during morning
and evening milking processes. Results of standard
chemical analysis for the examined milk components
are presented in Table 1. Collected samples varied
widely in milk composition and especially in fat content,
which tested the ability of NIR spectroscopy to provide
milk composition analysis at defined intervals in the
milking process.
The spectra of the milk samples taken during morn-
ing milking of one of the examined cows are shown in
Figure 1. The spectra were split into two regions at
1100 nm because of the change of detectors in NIRSys-
tem 6500 spectrophotometer, which were silicon for the
region from 700 to 1100 nm and lead sulfide (PbS) from
1100 to 2500 nm. Because of the water absorbance,
strong absorption bands around 960, 1440, and 1950
nm (21) dominated the spectra. The characteristic ab-
sorption bands of fat and other milk components such
as protein and lactose were very weak in comparison
with the water bands and were hard to detect. The
spectrum was shifting upwards with the fat content ele-
vation.
Preliminary investigations showed that smoothing
and pretreatment of milk spectral data were needed for
measurement of milk composition by NIRS. The best
calibration models for fat, total protein, and lactose
content determination, in terms of smoothing intervals
Journal of Dairy Science Vol. 82, No. 11, 1999
TSENKOVA ET AL.
and derivative windows, based on log (1/T) data, first
derivative, and second derivative transformation are
SD given in Tables 2 to 4. The comparison of the calibration
1.921 models was made on the basis of calibration and cross-
0.232 validation statistics that included standard error of cal-
0.163 ibration, coefficient of multiple correlations, SECV,
cross-validation correlation coefficient (CVr), and coef-
ficient of variation (CV) = SECV/Mean value) × 100.
We found that the accuracy of fat content determina-
tion of bovine milk depended strongly on the spectral
regions and path lengths. The best results were ob-
tained for the region from 1100 to 2400 nm with 1-mm
sample thickness (Table 2). The SECV for the model
based on the first derivative spectral data transforma-
tion with a 17-point derivative window was 0.110, the
CVr was 0.998, and the CV was 3.15%. This accuracy
of prediction was close to the accuracy of the reference
method. Figure 2 illustrates the relationship between
the chemical data for fat and the respective values pre-
dicted by the NIR calibration equation. The accuracy
of the model, based on log (1/T) spectral data, was simi-
lar to the best first derivative model.
In the spectral region from 700 to 1100 nm, the lowest
SECV and the highest CVr for fat content determina-
tion were obtained using spectral data for samples that
were 10 mm thick. The model derived using log (1/T)
data smoothed with windows with 13 data points had
SECV = 0.191, CVr = 0.995, and CV = 5.46%. The lowest
accuracy in fat determination was found for models
derived using spectral data from 700 to 1100 nm of
samples that were 1 mm thick.
These results were consistent with the investigation
of Pierce and Wehling (8) for determination of fat con-
Figure 1. Absorbance of near-infrared spectra: (log1/T) in the 700
to 2400 nm region of milk samples from one morning milking of cow
no. 987 (1, fat = 0.78%; 2, fat = 1.33%; 3, fat = 2.10%; 4, fat = 3.05%;
5, fat = 4.58%; and 6, fat = 6.48%).
 MILK COMPOSITION MEASUREMENT 2347
TABLE 2. Near-infrared spectra calibration and validation statistics for fat content determination in unho-
mogenized milk.
Smoothing Calibration statistics3
Spectral region interval or
and sample Spectral data1 derivative PLS2
thickness transformation windows factors SEC R SECV CVr CV%
700−1100 nm Log(1/T) 9 10 0.310 0.986 0.376 0.998 10.72
1 mm D1 9 9 0.284 0.988 0.347 0.981 9.91
D2 25 10 0.236 0.992 0.262 0.989 7.48
700−1100 nm Log(1/T) 9 10 0.172 0.996 0.213 0.994 6.09
4 mm D1 17 10 0.178 0.996 0.203 0.994 5.78
D2 25 8 0.180 0.996 0.203 0.994 5.78
700−1100 nm Log(1/T) 13 10 0.171 0.996 0.191 0.995 5.46
10 mm D1 13 10 0.167 0.996 0.193 0.994 5.52
D2 25 10 0.221 0.993 0.246 0.989 7.03
1100−2400 nm Log(1/T) 9 10 0.103 0.999 0.115 0.998 3.27
1 mm D1 17 10 0.098 0.999 0.110 0.998 3.15
D2 25 10 0.116 0.998 0.133 0.997 3.80
1
D1 = first derivative transformation, D2 = second derivative transformation, and T = transmittance.
2
Number of factors in the calibration model; PLS = partial least square.
3
SEC = Standard error of calibration, R = coefficient of multiple correlation, SECV = standard error of
cross-validation, CVr = cross-validation correlation coefficient, and CV = variation coefficient − (SECV/mean
value) × 100.

tent in Cheddar cheese. They found that the spectral
region from 1100 to 2500 nm performed better than did
the spectral region from 700 to 1100 nm.
For the 700 to 2400 nm region, SECV for lactose
content determination varied in a narrow range from
0.082 to 0.099, and the corresponding CV varied be-
tween 1.87 and 2.26% (Table 3). This accuracy of predic-
tion was close to the accuracy of the reference method.
The observed small total variation in the lactose content
explained the relatively low CVr that ranged from 0.773
to 0.827. The best calibration model for lactose content
determination was obtained when applied the first de-
rivative transformation of log (1/T) data in the 1100 to
2400 nm region with a 1-mm sample thickness. The
Figure 2. Correlation between measured and near-infrared spec-
troscopy (NIRS) predicted values for milk fat content (spectral region
from 1100 to 2400 nm; 1-mm sample thickness; first derivative
data transformation).
SECV for this model was 0.082, CVr was 0.827, and
the variation coefficient was 1.87%. Figure 3 illustrates
the relationship between chemical data for lactose and
the respective values predicted by the NIR calibration
equation. Accuracy of the model, based on the same
spectral region and log (1/T) data with a smoothing
interval of 17 data points, was similar. In the spectral
region from 700 to 1100 nm, the best model for lactose
content determination was obtained using log (1/T) data
with a 4-mm sample thickness, which had the lowest
SECV value of 0.089. This accuracy was very close to
the results in the region from 1100 to 2400 nm.
Table 4 shows that the accuracy of total protein con-
tent determination for bovine milk depended strongly
on the spectral region and the path length. The best
results were obtained for the region from 1100 to 2400
nm with a 1-mm sample thickness. The SECV for the
best model, based on the first derivative spectral data
transformation with a 25-point derivative window, was
0.096, CVr was 0.848, and CV was 3.06%. This result
showed that the accuracy of this method was close to
the accuracy of the reference method. Figure 4 illus-
trates the relationship between the data for chemical
analysis of total protein and the respective values pre-
dicted by the NIR calibration equation. The SECV val-
ues for the best models, based on log (1/T) and second
derivative transformation of the spectral data, 0.115
and 0.118 respectively, were similar to the SECV of the
first derivative model.
Satisfactory accuracy for determination of total pro-
tein content in the near NIR region, from 700 to 1100
nm, was found when a 1-mm sample thickness was
used. The lowest SECV and the highest CVr were ob-

Journal of Dairy Science Vol. 82, No. 11, 1999

2348 TSENKOVA ET AL.

TABLE 3. Near-infrared spectra calibration and validation statistics for lactose content determination in
unhomogenized milk.
Smoothing Calibration statistics3
Spectral region interval or
and sample Spectral data1 derivative PLS2
thickness transformation windows factors SEC R SECV CVr CV%
700−1100 nm Log(1/T) 13 7 0.086 0.813 0.092 0.778 2.09
1 mm D1 13 5 0.087 0.808 0.092 0.779 2.08
D2 25 6 0.085 0.816 0.092 0.800 2.08
700−1100 nm Log(1/T) 21 10 0.082 0.829 0.089 0.784 2.02
4 mm D1 13 8 0.083 0.822 0.090 0.776 2.05
D2 25 9 0.084 0.818 0.089 0.781 2.03
700−1100 nm Log(1/T) 5 6 0.090 0.840 0.096 0.809 2.18
10 mm D1 25 6 0.091 0.836 0.094 0.820 2.13
D2 25 6 0.091 0.838 0.099 0.795 2.26
1100 2400 nm Log(1/T) 17 10 0.077 0.857 0.086 0.773 1.95
1 mm D1 25 10 0.076 0.854 0.082 0.828 1.87
D2 25 10 0.081 0.841 0.091 0.783 2.07
1
D1 = first derivative transformation, D2 = second derivative transformation, and T = transmittance.
2
Number of factors in the calibration model, PLS = partial least square.
3
SEC = Standard error of calibration, R = coefficient of multiple correlation, SECV = standard error of
cross-validation, CVr = cross-validation correlation coefficient, CV = variation coefficient − (SECV/mean
value) × 100.

tained using log (1/T) data, smoothed with a 9 data
points window: SECV = 0.129, CVr = 0.706, and CV =
4.13%. The results for total protein determination using
this sample thickness were very close to the results
when the region from 1100 to 2400 nm was used. The
lowest accuracy in total protein determination was
found for models that were derived using spectral data
from the 700 to 1100 nm and combined with a 10-mm
sample thickness. This accuracy was inadequate for
practical applications.
No particular spectral data treatment proved to be
the best for the tested combinations of spectral regions
Figure 3. Correlation between measured and near-infrared spec-
troscopy (NIRS) predicted values for lactose content of milk (spectral
region from 1100 to 2400 nm; 1-mm sample thickness; first derivative
data transformation).
and sample thicknesses. Different smoothing intervals
or different derivative windows gave different accura-
cies for fat, total protein, and lactose determinations.
Models based on the second derivative data transforma-
tion tended to yield better results when using larger
window sizes. The influence of the data transformation
on the accuracy of fat and protein content determina-
tions was smaller than the influence of the spectral
region and sample thickness.
The PLS modeling aided not only the development
of quantitative models for milk composition determina-
tion but was also used as a tool for discerning the loca-
tion of information related to fat, total protein, and
lactose. No information was found in the literature con-
cerning determination of milk composition in the near
NIR region. In this study, regression vectors of the cali-
bration models were studied, based on log (1/T) spectral
data in this region. High positive coefficients in the
regression vector for the model of fat determination in
the 700 to 1100 nm spectral region with a 10-mm sam-
ple thickness were found at 730, 776, 930, 968, 990,
1026, 1076, and 1092 nm (Figure 5). The highest posi-
tive coefficient was observed at 930 nm. The absorption
at 930 nm could be associated with the C-O band of oil.
The regression vector for the best lactose determina-
tion model in the region from 700 to 1100 nm with a
4-mm sample thickness is presented in Figure 6. High
positive coefficients in the regression vector were found
at 734, 750, 786, 812, 908, 974, 982, and 1064 nm. The
highest positive coefficient was observed at 1064 nm.
High positive coefficients in the regression vector for
the total protein determination model in the 700 to

Journal of Dairy Science Vol. 82, No. 11, 1999

 MILK COMPOSITION MEASUREMENT 2349
TABLE 4. Near-infrared spectra calibration and validation statistics for total protein content determination
in unhomogenized milk.
Smoothing Calibration statistics3
Spectral region interval or
and sample Spectral data1 derivative PLS2
thickness transformation windows factors SEC R SECV CVr CV%
700−1100 nm Log(1/T) 9 10 0.112 0.792 0.129 0.706 4.13
1 mm D1 9 10 0.117 0.771 0.134 0.676 4.27
D2 13 10 0.115 0.782 0.134 0.676 4.27
700−1100 nm Log(1/T) 9 10 0.127 0.714 0.148 0.571 4.74
4 mm D1 9 10 0.129 0.704 0.150 0.556 4.79
D2 25 10 0.138 0.650 0.151 0.539 4.82
700−1100 nm Log(1/T) 9 7 0.175 0.602 0.191 0.482 6.11
10 mm D1 17 10 0.173 0.620 0.191 0.483 6.10
D2 25 10 0.184 0.552 0.202 0.386 6.46
1100−2400 nm Log(1/T) 17 10 0.100 0.840 0.115 0.774 3.67
1 mm D1 25 10 0.085 0.886 0.096 0.848 3.06
D2 25 10 0.104 0.826 0.118 0.760 3.77
1
D1 = first derivative transformation, D2 = second derivative transformation, and T = transmittance.
2
Number of factors in the calibration model, PLS = partial least square.
3
SEC = Standard error of calibration, R = coefficient of multiple correlation, SECV = Standard error of
cross-validation, CVr = Cross-validation correlation coefficient, CV% = variation coefficient − (SECV/mean
value) × 100.

1100 nm region with a 1-mm sample thickness were
found at 726, 736, 760, 776, 794, 880, 902, 952, and
1034 nm (Figure 7). The absorptions at 760, 880, and
902 nm could be associated with the N-H third overtone
bands of protein and at 1034 nm with the N-H second
overtone band. The peak at 952 nm was very close to
the water band and probably was connected with the
influences of water-soluble protein of milk on water ab-
sorption.
CONCLUSIONS
The NIR spectroscopy was an adequate method for
determination of fat, total protein, and lactose of unho-
mogenized cow milk for wide range of milk composition
changes, which were observed in the milking process
and over one lactation.
Spectral region and sample thickness were found to
be significant factors for determination of milk fat and
total protein. The best accuracies for fat, total protein,
and lactose content determinations were obtained with
the 1100 to 2400 nm region, combined with a 1-mm
sample thickness and first derivative data transforma-
tion. For the spectral region from 700 to 1100 nm, the
best results for fat measurement were obtained for sam-
ples that were 10 mm thick; total protein required a 1-
mm sample thickness. The different sample thicknesses
in the region from 700 to 1100 nm did not have strong

Figure 4. Correlation between measured and near-infrared spec-

troscopy (NIRS) predicted values for total protein content of milk Figure 5. Regression vector plot of the model for milk fat determina-
(spectral region from 1100 to 2400 nm; 1-mm sample thickness; first tion (spectral region from 700 to 1100 nm; 10-mm sample thickness;
derivative data transformation). log (1/T) data).

Journal of Dairy Science Vol. 82, No. 11, 1999

2350
Figure 6. Regression vector plot of the model for milk lactose deter-
mination (spectral region from 700 to 1100 nm; 10-mm sample thick-
ness; log (1/T) data).
influence on the accuracy of lactose content determina-
tion. The influence of spectral data treatment on the
accuracy of determining milk fat and total protein con-
tents was less than the influence of the spectral region
and the sample thickness. The best results for determi-
nation of fat, total protein, and lactose contents in the
region from 700 to 1100 nm had accuracies very close
to that obtained in the region from 1100 to 2400 nm.
Our study showed that the short-wave NIR region could
be successfully used for on-line measurement of milk
composition in the process of milking for different cows
and different stages of their lactation.
The wavelengths with the largest contribution for
determination of constituents of milk in the short-wave
NIR spectral region from 700 to 1100 nm were found
to be 730, 776, 930, 968, 990, 1026, 1076, and 1092 nm
for fat content; 734, 750, 786, 812, 908, 974, 982 and
Figure 7. Regression vector plot of the model for milk total protein
determination (spectral region from 700 to 1100 nm; 1-mm sample
thickness; log (1/T) data).
Journal of Dairy Science Vol. 82, No. 11, 1999
TSENKOVA ET AL.
1064 nm for lactose content; and 726, 736, 760, 776, 794,
880, 902, 952, and 1034 nm for total protein content.
ACKNOWLEDGMENTS
This work was supported by the Program for Promo-
tion of Basic Research Activities for Innovative Biosci-
ence (PROBRAIN), Ministry of Agriculture, Fisheries
and Food, Japan. We thank the staff of the University
Farm at Hokkaido University, Sapporo, Japan for pro-
viding cows for the experiment. The authors are grate-
ful to GL Science Inc., Tokyo, Japan for providing the
software Pirouette, Version 2.0.
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