Meat and milk compositions of bovine clones

X. Cindy Tian, Chikara Kubota, Kunihito Sakashita, Yoshiaki Izaike, Ryoichi Okano, Norio
Tabara, Carol Curchoe, Lavina Jacob, Yuqin Zhang, Sadie Smith, Charles Bormann, Jie Xu,
Masumi Sato, Sheila Andrew, and Xiangzhong Yang
PNAS 2005;102;6261-6266; originally published online Apr 13, 2005;
doi:10.1073/pnas.0500140102
This information is current as of April 2007.

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Notes:
Meat and milk compositions of bovine clones
X. Cindy Tian*†, Chikara Kubota†‡, Kunihito Sakashita§, Yoshiaki Izaike¶, Ryoichi Okano§, Norio Tabara§,
Carol Curchoe*, Lavina Jacob*, Yuqin Zhang*, Sadie Smith*, Charles Bormann*, Jie Xu*, Masumi Sato储,
Sheila Andrew*, and Xiangzhong Yang*,**
*Center for Regenerative Biology兾Department of Animal Science, University of Connecticut, Storrs, CT 06269; ‡Kagoshima Prefectural Cattle Breeding
Development Institute, 2200 Tsukino Osumi So-Gun Kagoshima 899-8212, Japan; §Kagoshima Prefectural Livestock Experiment Station, 2440 Uenodan
Kokubu, Kagoshima 899-4461, Japan; ¶Department of Developmental Biology, National Institute of Agrobiological Science, 2 Ikenodai, Tsukuba,
Ibaraki 305-8602, Japan; and 储Comparative Pathology Section, Kyushu Research Station, National Institute of Animal Health, 2702 Chuzan
Kagoshima, Kagoshima 891-0105, Japan

Edited by R. Michael Roberts, University of Missouri, Columbia, MO, and approved March 16, 2005 (received for review January 7, 2005)

The technology is now available for commercial cloning of farm breed-matched comparator animals from conventional reproduc-
animals for food production, but is the food safe for consumers? tion. All of the experimental animals used for the comparisons were
Here, we provide data on >100 parameters that compare the managed under the same conditions and received the same diet.
composition of meat and milk from beef and dairy cattle derived This report addresses the scientific and public concerns of the
from cloning to those of genetic- and breed-matched control physiology and safety of the meat and milk products from beef and
animals from conventional reproduction. The cloned animals and dairy animal clones.
the comparators were managed under the same conditions and
received the same diet. The composition of the meat and milk from Methods
the clones were largely not statistically different from those of Cloned Beef and Dairy Cattle. Our beef and dairy animal clones were
matched comparators, and all parameters examined were within produced by somatic cell nuclear transfer by using cultured skin
the normal industry standards or previously reported values. The fibroblasts or cumulus cells from an adult Japanese Black beef bull
data generated from our match-controlled experiments provide or a Holstein dairy cow (Fig. 1). Our beef clones were produced in
science-based information desired by regulatory agencies to ad- 1998; male clones of a farm animal species had not been produced
dress public concerns about the safety of meat and milk from previously. The donor bull (Kamitakafuku), a superior breeding
somatic animal clones. stud bull with superior marbling traits, was 17 years old when we
produced six bull clones of him (5). Four clones survived normally,
cloned cattle 兩 food safety 兩 clone health and two of these clones were selected randomly for serial cloning,
semen quality, and breeding performance analyses (6). The other
two beef clones were slaughtered and subjected to standard meat
S omatic cell cloning by nuclear transfer has potential agricultural
applications for duplicating food animals with desired genetic
merit. However, somatic cloned animals have been associated with
analyses in this study. We produced 10 diary clones from skin
fibroblast (n ⫽ 4) and cumulus cells (n ⫽ 6) of a Holstein cow at
aberrant gene expression (1–3), as well as developmental abnor- 13 years of age, between June and August 1999 (7). Four of these
malities and high neonatal death rates. These findings suggest the clones, all derived from cumulus cells, survived and are healthy. We
incomplete reactivation of some inactivated genes from the differ- have studied their telomere lengths (7), expression of X-linked
entiated somatic donor cells. Because of limited knowledge of the genes (1), onset of puberty (8), growth endocrinology (9), and
behavior (10). All animal use was approved by the institutional

AGRICULTURAL
nature of gene dysregulation in clones, public debate has arisen as

SCIENCES
to whether food products from animal clones are safe for human animal care and use committees at the University of Connecticut
consumption. In the United States, the Food and Drug Adminis- (dairy) or the Kagoshima Prefectural Institute of Cattle Breeding
tration’s Center for Veterinary Medicine has asked companies not and Development (beef).
to introduce animal clones, their progeny, or their food products,
such as milk or meat, into the human or animal food supply Comparison of Milk Production. In the present study, the four live
(www.fda.gov兾cvm兾index兾updates兾clones.htm). The U.S. Food dairy clones and four age and parity-matched comparator heifers
and Drug Administration requested that producers abstain from from natural reproduction were raised in the same facility from 2
placing edible products from clones into the food supply until the months of age. All animals were subjected to the same diet and
agency considers the safety of their products based on scientific management protocols and were bred by artificial insemination or
information gained from the direct evaluation of safety. To date, no natural breeding starting at 14–15 months of age. Immediately after
animals cloned from somatic nuclei, or their products, have been calving, we monitored milk production by collecting samples three
permitted to enter the food chain in any country (4). Information times daily during the entire first lactation. The total amount of milk
on the composition of meat and milk from somatic clones of food produced in the first 305 days of lactation, the standard lactation
animals is extremely limited and highly desired by federal regulatory period in dairy cattle, was compared among the clones, to their
agencies concerned with food safety. Commissioned by the U.S. matched comparators and to the production records of the clones’
Food and Drug Administration, the National Academy of Sciences genetic donor cow.
was charged to identify any safety concerns that animal clones might
present to humans, animals, and the environment. The National Comparison of Milk Composition. To compare the milk composi-
Academy of Sciences report concludes that clones are not likely to tions, two milk samples were collected from each of the three
pose a food consumption risk, but the National Academy of milkings on a given day of each week, throughout the entire first
Sciences states that information on compositions of the products of lactation. One of these milk samples was delivered to Dairy One
animal clones is needed to decrease food safety uncertainties
(www.nap.edu兾catalog兾10418.html). Thus, we have conducted ex-
This paper was submitted directly (Track II) to the PNAS office.
tensive comparisons of the composition of milk and meat from
somatic cloned animals to those from naturally reproduced com- Abbreviation: DHIA, Dairy Herd Improvement Association.

parator animals. Here, we provide data on ⬎100 parameters that †X.C.T. and C.K. contributed equally to this work.
compare the composition of meat and milk from beef and dairy **To whom correspondence should be addressed. E-mail: xiangzhong.yang@uconn.edu.
cattle derived from somatic cloning to those of genetic- and © 2005 by The National Academy of Sciences of the USA

www.pnas.org兾cgi兾doi兾10.1073兾pnas.0500140102 PNAS 兩 May 3, 2005 兩 vol. 102 兩 no. 18 兩 6261– 6266

 Fig. 2. Analyses of milk production and compositions of somatic clones and
Fig. 1. Somatic cloning and clones. (A) Source and production procedure of
matched comparator cows. (A) Representative first lactation curves of a clone
the cloned dairy and beef cattle used for this study. (B) The genetic relation-
and a comparator cow. (B) Milk total protein (percentage), total fat (percent-
ship among the beef clones and their comparators. Squares indicate males,
age), lactose (percentage), total solids (percentage), milk urea nitrogen (mg兾
and circles indicate females. The same colors indicate identical genetic
dl) and somatic cell count (⫻ 103兾ml). (C) A representative image of protein
makeup. The genetic comparators share 25% of genetic identity with the
profile analysis of milk samples from clones (lanes 1–3) and comparator cows
clones, whereas the breed comparators are not related genetically to the
(lanes 4 –7) by SDS兾PAGE. Lane 8, molecular mass markers. The four major
clones or the donor animal.
bands are (from top to bottom identified on the basis on their molecular
mass): ␣-caseins, ␤-caseins, ␬-caseins, and ␤-lactoglobulins.

Cooperative (Ithaca, NY), a Dairy Herd Improvement Association

(DHIA)-designated laboratory, for analyses of total protein (per- facility and subjected to the same diet and management. The
centage), total fat (percentage), lactose (percentage), total solids genetic matched comparator bulls were produced by artificial
(percentage), milk urea nitrogen (mg兾dl), and somatic cell counts
insemination using semen from the son of the original donor bull
(⫻ 103 per ml). All these parameters are routinely monitored for
(Kamitakafuku), and thus, they are genetic ‘‘nephews’’ and share
the dairy industry by the DHIA (www.dhia.org). We used the
second set of the milk samples (frozen at ⫺20°C) for monthly 25% of their genetic makeup with the clones (Fig. 1B). Additionally,
analyses of protein profiles by denaturing SDS兾PAGE stained with 20 mature Japanese Black beef cattle (referred to as breed com-
Coomassie blue. The gel images were scanned, and the relative parators), at the Kagoshima Prefectural Livestock Station or Japan
quantities of each band were determined by using the QUANTITY Meat Grading Association, were also used to establish the normal
ONE software program (Bio-Rad). Additionally, we also measured range for each parameter analyzed. All bulls in this study were
antobody concentrations (IgM, IgA, and IgG) in the colostrum castrated at 3 months of age and were then given a standard growing
from the first milking by using the Single Radial ImmunoDiffusion ration from 8 to 26 months of age, according to the normal practice
kit (VMRD, Pullman, WA). for the beef breed in Japan. The animals were then slaughtered and
examined to determine meat quality and carcass composition by
Comparison of Meat Composition. We compared our two beef clones using beef industry standard protocols (www.acess.gpo.gov兾nara兾
with eight genetically matched comparators by using the same cfr兾waisidx㛭99兾9cfrv2㛭99.html). The following parameters were
analyses protocols (Fig. 1B). All animals were raised in the same analyzed and compared between the beef clones and their genetic

6262 兩 www.pnas.org兾cgi兾doi兾10.1073兾pnas.0500140102 Tian et al.

 Table 1. Parameters of the left dressed carcass
Genetic Breed
Clone comparator comparator
Parameter (n ⫽ 2) (n ⫽ 8) (n ⫽ 20)

Carcass weight, kg 425.5 ⫾ 16.8 457.4 ⫾ 30.4 457.0 ⫾ 43.7

Rib eye area, cm2 57.0 ⫾ 1.41 58.9 ⫾ 5.1 55.4 ⫾ 6.4
Rib thickness, cm 8.7 ⫾ 0.00 7.8 ⫾ 0.6 7.6 ⫾ 0.6
s.c. fat thickness, cm 3.3 ⫾ 0.00 3.1 ⫾ 0.4 3.3 ⫾ 0.5
Yield score, % 74.4 ⫾ 0.00* 73.8 ⫾ 0.3 73.0 ⫾ 0.8
Beef marbling standard 8.0 ⫾ 1.41 6.5 ⫾ 0.9 5.2 ⫾ 1.5
Beef color standard 3.0 ⫾ 0.00 2.9 ⫾ 0.4 3.3 ⫾ 0.7
Beef fat standard 3.0 ⫾ 0.00 2.9 ⫾ 0.4 3.1 ⫾ 0.5

Results are presented as mean ⫾ SD. *, Significant difference was detected

between clones and comparators.

acid, stearic acid, oleic acid, linoleic acid, and linolenic acid) of five
major fat tissues (s.c. fat, intra- and inter-muscular fats, celom fat,
and kidney leaf fat); analyzed by gas chromatography after lipid
extraction at the Japan Food Research Laboratories; (vi) amino
acid composition of the longissimus thoracis muscle; determined by
an amino acid analysis system (Shimadzu) at Kagoshima Prefectural
Livestock Station (three controls were used for this analysis); and
(vii) histopathology of all organs, examined at the National Institute
of Animal Health, Kyusyu, Japan.

AGRICULTURAL
SCIENCES
Fig. 3. Body organ parameters for clones (n ⫽ 2), genetic comparators (n ⫽
8, except in A where n ⫽ 3), and breed comparators (n ⫽ 20). (A) Proportions
(percentage; means ⫾ SD) of organ or body part (g) over body weight (kg). (B)
The proportions over body weight of various muscles or fat tissues (percent-
age; means ⫾ SD). *, Significant difference was detected between clones and
comparators.

and breed comparators: (i) organ or body part weights; (ii) total
proportion of meat and fat in the dressed carcass; (iii) cross section
of the left dressed carcass between the sixth and seventh rib,
following the standard methods of the Japan Meat Grading Asso-
ciation (11); (iv) the moisture, crude protein, and crude fat contents
of six muscles (infraspinatus, longissimus thoracis, latissimus dorsi,
Fig. 4. Parameters (percentage; means ⫾ SD) for clones (n ⫽ 2), genetic
adductor, biceps femoris, and semitendinosus); measured by the comparators (n ⫽ 8), and breed comparators (n ⫽ 20). (A) Muscle moisture. (B)
Kjeldahl analysis method from the Official Methods of Analysis of Muscle crude fat. (C) Muscle crude protein. (D) Amino acid composition of
AOAC International by the Soxtec method (12); (v) fatty acid longissimus thoracis muscle (mg兾100 g of muscle. Results are means ⫾ SD; *,
composition (lauric acid, myristic acid, palmitic acid, palmitoleic significant difference was detected between clones and comparators.

Tian et al. PNAS 兩 May 3, 2005 兩 vol. 102 兩 no. 18 兩 6263

Fig. 5. Comparisons of fatty acid composition (g兾100 g) among clones (n ⫽ 2), genetic comparators (n ⫽ 8), and breed comparators (n ⫽ 20) in various tissues.
(A) Subcutaneous fat. (B) Kidney leaf fat. (C) Intramuscular fat in longissimus thoracis muscle. (D) Intramuscular fat in semitendinosus muscle. (E) Intermuscular
fat. Results are means ⫾ SD; *, significant difference was detected between clones and comparators.

Data Analyses. Milk production and composition data were sub- analysis is shown in Fig. 2C. Four major bands, ranging from ⬇17
jected to a mixed model analysis by using the General Linear Model to 35 kDa and representing ␣-caseins, ␤-caseins, ␬-caseins, and
(SAS 9.0, SAS Institute, Cary, NC) with week as a repeated measure. ␤-lactoglobulins, were observed to be consistent in all milk samples
The somatic cell count data were analyzed after a log transforma- from either the clones or their comparators. Minor bands were also
tion. Data presented in figures are least-square means. We com- present at high and low molecular masses in all samples, indistin-
pared 90% confidence intervals of each parameter of meat com- guishable among clones and comparators. There was no significant
position in a pairwise manner to determine any significant difference in the percentages of each major constituent protein
difference of biological relevance. between milk samples from the clones and their comparators.
Antibodies in the colostrum from the clones ranged from 2,000 to
Results 15,000 mg兾dl, 70 to 360 mg兾dl, and 125 to 500 mg兾dl for IgG, IgA,
Milk Production. All cloned and comparator animals were bred with and IgM, respectively. These values for antibody concentrations
semen from different bulls and delivered normal calves during three were in the typical range for antibody composition of colostrums
consecutive parturitions at the expected due dates, with the excep- (14), and the concentrations of antibodies in colostrum from the
tion of one parturition. The exception was that one of the clones clones and their comparators appeared similar. These results indi-
(clone B) gave birth to a stillborn calf, 2 weeks prematurely at her cate that the quality of colostrum from clones is sufficient for the
first parturition, and did not have full udder development at the nutritional and health requirements of their calves.
commencement of lactation. All of the other pregnancies produced
normal calves in terms of their gestation lengths, ease of calving, Meat Composition. We analyzed ⬎100 parameters concerning the
and birth weights. Together, we collected ⬎1,000 milk samples, and quality of meat from our beef clones and matched comparator
the representative production curves of a clone and a comparator animals, using the standard analysis methods of the industry. The
animal are shown in Fig. 2A. All of the clones and their matched 90% confidence intervals (C.I.) of each parameter of meat
comparators showed similar, normal lactation curves (13); milk composition in a pairwise manner were used to determine any
production increased during the first month of lactation and then significant difference of biological relevance. Overlap of C.I.s for
declined progressively for the remainder of the lactation period. each paired comparison (clone vs. genetic comparator and clone
The amount of milk produced by the four clones (8,646.1 ⫾ 743.8 vs. breed comparator) were not significantly different. Our
kg) in the first lactation was not significantly different compared results indicated that no significant difference was detected in
with that of matched comparator cows (9,507.8 ⫾ 743.8 kg). Clone ⬎90% of all parameters examined (Figs. 3–5 and Table 1). There
B, who gave birth prematurely, produced 30% less milk (6,339.3 kg) were, however, 12 instances in which the clones and genetic
than the average of the other three clones (9,378.4 kg). The reason comparators showed differences, and these were as follows: the
that the donor cow was cloned was that she held one of the highest amount of mesentery fat (Fig. 3A); yield score (Table 1); the
production records in the herd in her best lactation period (15,875.9 proportion of longissimus thoracis muscle over body weight (Fig.
kg). As expected, the production of the three clones in their first 3B); the muscle moisture (Fig. 4A) and the amount of crude
lactation (9,378.4 kg) was similar to that of the donor animal in her protein in the semitendinosus muscle (Fig. 4C); the amount of
first lactation (8,990.7 kg; P ⬎ 0.05). linolenic acid in kidney leaf fat (Fig. 5B) in the longissimus
thoracis (Fig. 5C) and semitendinosus (Fig. 5D) muscles; and the
Milk Composition. No significant difference was detected between amount of oleic acid, palmitic acid, palmitoleic acid, and linoleic
the composition of milk from the clones and the matched compar- acid in the semitendinosus muscle (Fig. 5D). All these param-
ator cows (Fig. 2B). A representative image of the protein profile eters concerning the amount of mesentery fat and fatty acids in

6264 兩 www.pnas.org兾cgi兾doi兾10.1073兾pnas.0500140102 Tian et al.

 Table 2. Pathological observations on the clones
Clone

Organ Examination level Observation 1 2

Liver Gross appearance Normal Normal

Histological findings Irregular arrangement of hepatic ⫺ ⫺
cords
Focal necrosis ⫺ ⫺
Inflammation ⫺ ⫺
Fibrosis ⫺ ⫺
Kidney Gross appearance Structure Normal Normal
Urinary calculus (collecting duct) ⫹* ⫹*
Histological findings
Cortex
Glomerulus Varying size ⫺ ⫺
Immature form ⫺ ⫺
Proliferation of the mesangial cells ⫺ ⫺
Thickened basement membrane ⫺ ⫺
Renal tubules Cystic tubules ⫺ ⫺
Urinary casts ⫺ ⫺
Atrophied or immature tubules ⫺ ⫺
Thickened basement membrane ⫺ ⫺
Abnormal cell infiltration ⫺ ⫺
Medulla Urinary cast ⫺ ⫺
Cystic tubule ⫺ ⫺
Inflammation ⫺ ⫺
Lung Gross appearance Normal Normal
Histological findings Atelectasis ⫺ ⫺
Inflammation ⫺ ⫺
Heart Gross appearance Normal Normal
Histological findings
Cardiac muscle cells Irregular arrangement ⫺ ⫺
Necrosis ⫺ ⫺
Inflammation ⫺ ⫺
Spleen Gross appearance Normal Normal
Histological findings
Inflammation ⫺ ⫺
Hyperplasia ⫺ ⫺
Adrenal gland Gross appearance Normal Normal

AGRICULTURAL
⫺ ⫺

SCIENCES
Histological findings Irregular arrangement of cortex cells
Hyperplasia ⫺ ⫺
Thyroid Gross appearance Normal Normal
Histological findings Abnormal follicles ⫺ ⫺
Abnormal follicular epithelium ⫺ ⫺

*, Significant difference was detected between clones and comparators.

the meat兾fat were significantly higher in the clones than in their reproduced comparator animals, using standard protocols well
genetic or breed comparators, except for crude protein or muscle established in the beef and dairy industries. We found no significant
moisture in semitendinosus muscle. differences in the composition of milk from cloned animals com-
To determine the comparative health and pathology of all organs pared with the comparator animals managed under the same
from the two clones used for meat analyses, the organs were conditions. Our results of the milk analyses using the DHIA
subjected to histological analyses after slaughter. Both clones were standards suggest that healthy clones not only are normal them-
normal in all their organs, including liver, kidney, lung, heart, selves based on previously examined parameters, such as telomere
spleen, and the adrenal and thyroid glands (Table 2). No macro- lengths (7), onset of puberty (8), reproduction and lactation (17,
scopical or microscopical abnormalities were observed in the 18), growth endocrinology (9), expression of X-linked genes (1),
and behavior (10) but also appear to have normal gene expression
clones, except for the kidney urinary calculi (collecting ducts).
in their mammary tissues. This normality is because the production
However, these calculi are often detected in the usual beef cattle of each milk protein constituent involves the elaborate regulatory
because of a feeding peculiarity (15). function of many proteins and enzymes, and any abnormal gene
expression would likely be reflected by imbalances in the constit-
Discussion
uents of the milk. Furthermore, our finding that there were no
In a recent report, the composition of milk from somatic cow clones differences in somatic cell counts, which is a parameter used to
was analyzed, but the findings were confounded with different diets detect subclinical mastitis, demonstrates that these clones were not
and management (16), which are known to affect milk production more susceptible than the comparator animals to this mammary
and composition. In the present study, we compared the compo- gland disease that is associated with lactation.
sition of meat and milk from our somatic beef (5) and dairy (7) For the milk production comparison, we found that all clones and
cattle clones to those of age-, genetic-, and breed-matched naturally their matched comparators showed similar and normal lactation

Tian et al. PNAS 兩 May 3, 2005 兩 vol. 102 兩 no. 18 兩 6265

curves (13). The slight improvement in the milk production of the range of the previously recorded industry standards (19, 21, 22).
clones over the donor cow at their first lactation was likely due to Therefore, none of these parameters are of public concern.
improved nutrition and management practices that have evolved in To our knowledge, there have been no previous reports on the
the last 13 years in the dairy industry (www.usda.gov兾nass兾pubs兾 organ histology, composition, and quality of meat from somatic
histdata.htm). Overall, our data on comparing various aspects of cloned animals for potential human consumption. Previously,
milk, including protein profiles, antibody levels, composition, and there has been one study evaluating the meat of animals cloned
production from somatic cloned animals with naturally reproduced from embryonic cells (23). Those results, however, do not
comparator animals are very comprehensive and convincing. provide insight into the products from animals cloned from adult
One of our reasons for cloning the Japanese Black beef breeding somatic cells and could not be fully justified to serve as the
bull was his top-ranking breeding value due to an excellent meat scientific basis to address public concerns on the food safety of
somatic animal clones. This is because embryonic animal clones
marbling score. The popularity of this donor bull was shown by the
are produced from blastomeres of fertilized embryos at a very
fact that ⬎350,000 cows were inseminated with the bull’s semen and
early stage of development, and thus, embryonic clones may
that he was the sire of ⬎165,000 offspring at the time we cloned him undergo little gene reprogramming during their development.
in 1998. The two clones of this bull were found to have an average This is likely why food products from embryonic animal clones
marbling score of 8 of 12, whereas the average score for the breed have been used for human consumption and their safety has not
is only 5.2. The genetic comparators in this study had a marbling been a public concern.
score of 6.5, which is between the clones and the breed average In summary, we conclude that most parameters of the com-
(Table 1). This score is what might be expected, considering the fact position of the meat and milk from somatic animal clones were
that the genetic comparators were also descendents of and had not significantly different from those of their genetically
some genetic influence (25%) from the donor bull. Furthermore, matched comparators or industry breed comparators, and that
the two clones had nearly identical marbling patterns of the muscles all parameters examined in this study were within the normal
imaged at the sixth and seventh rib (photos not shown), using the range of beef and dairy products approved for human consump-
standard comparison for this breed, suggesting a strong genetic tion. It is important to note that this study was conducted with
influence on this production trait in the Japanese Black Beef cattle. a relatively small number of diary and beef clones, and the clones
In the present study, we analyzed ⬎100 parameters concerning of each breed were derived from a single genetic source. The
the quality of meat from our beef clones, and the prevailing experiments presented here, however, are a pilot study to
majority of these parameters did not differ from those of the provide guidelines for more conclusive studies with larger num-
matched comparator animals. Among the 12 parameters differ- bers of clones from different genetic backgrounds, to further
entially detected between the clones and comparator animals, 8 increase the consumers’ confidence concerning product safety of
were related to the amount of fat or fatty acids in the meat兾fat somatic cloned food animals.
(high levels in clones). Animals with more fat or fatty acids in
We thank K. Kouri, C. Moyes, D. Yang, P. Amond, K. Kawabata, M.
meat兾fat are more valuable in Japanese Black beef and have Ozono, M. Yonemaru, and the farm crew at the Kellogg Dairy Center
been selected for. The fact that both clones had consistently at the University of Connecticut for help with milk sample collection
higher amounts of mesentery fat and fatty acids compared with and兾or shipping; the farm crew at the Kagoshima Prefectural Livestock
the comparators is hardly surprising because these two clones are Experiment Station for taking care of cloned bulls; T. Onitsuka for
genetic copies of a top breeding bull and they both exhibited the histopathological analyses; T. Koga, Y. Da, L. Kuo, J. Riesen, and T.
most preferable values as expected (19, 20). The other four Hoagland for help with statistical analyses; and C. Faustman, R. Foote,
and M. Julian for critical reading of the manuscript. This work was
parameters found different between clones and comparators: supported by grants from the U.S. Department of Agriculture and
yield score, the proportion of longissimus thoracis muscle to Connecticut Innovations, Inc. (to X.Y. and X.C.T.); and from the
body weight, the muscle moisture, and the amount of crude Kagoshima Prefecture and the National Institute of Agrobiological
protein in the semitendinosus muscle, all fall within the normal Science (to C.K.).

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