Composition of milk from minor dairy animals and buffalo breeds: a biodiversity perspective

ReviewReceived: 3 May 2011 Revised: 13 August 2011 Accepted: 10 September 2011 Published online in Wiley Online Library:

(wileyonlinelibrary.com) DOI 10.1002/jsfa.4690

Composition of milk from minor dairy animalsand buffalo breeds: a biodiversity perspectiveElinor Medhammar, Ramani Wijesinha-Bettoni, Barbara Stadlmayr,Emma Nilsson, Ute Ruth Charrondiere∗ and Barbara Burlingame

Abstract

A comprehensive review is presented of the nutrient composition for buffalo, mare, and dromedary camel milks at thelevel of breed, and species-level data for yak, mithun, musk ox, donkey, Bactrian camel, llama, alpaca, reindeer and moosemilks. Average values of nutrients were calculated and compared. Interspecies values (g 100 g−1) were 0.7–16.1 for total fat,1.6–10.5 for protein, 2.6–6.6 for lactose, and 67.9–90.8 for water. Reindeer and moose milks had the highest fat and proteinconcentrations and the lowest lactose contents. Mare and donkey milks had the lowest protein and fat contents, in additionto showing the most appropriate fatty acid profile for human nutrition. Dromedary camel milk was most similar to cow milk inproximate composition. Moose milk was the richest in minerals, having values as high as 358 mg 100 g−1 for calcium, 158 mg100 g−1 for sodium and 150 mg 100 g−1 for phosphorus. Interbreed differences of 4 g 100 g−1 were observed in total fat inbuffalo, yak, mare and dromedary camel milks. Large interbreed differences were also present in the mineral contents in mare,buffalo and dromedary camel milks. By bringing together these compositional data, we hope to usefully widen the biodiversityknowledge base, which may contribute to the conservation and sustainable use of milk from underutilized dairy breeds andspecies, and to improved food and nutrition security, particularly in developing countries.c© 2011 Society of Chemical Industry

Keywords: milk; biodiversity; nutrient; composition; breed; species

INTRODUCTIONMilk is one of the most nutritionally complete foods. It containsall the nutrients necessary to sustain life of the neonate and theyoung infant, and adds quality to the human diet. Milk is anexcellent source of macro- and micro nutrients, and thereforecan play an important role in helping individuals to meet theirnutritional requirements.1 Milk protein contains all the essentialamino acids and thus provides an important source of proteinof high biological value, especially useful in developing countrieswhere rice or tubers are staples.2 Dairy animals are a key factorin household food security for small-scale livestock holders, whosupply the vast majority of milk in developing countries.3 Cow,goat and sheep milks account for about 87% of the global milkproduction.4 However, minor dairy animal species are nutritionallyand economically important in several countries.

The significance of biodiversity, defined as variation in foodcomposition at the species level and below, has recently beendiscussed in depth.5,6 The general global trend has beentowards diet simplification, with negative impact of food security,nutritional balance and the growing incidence of chronic diseases.Biodiversity contributes to a more diverse diet that can reversethis unhealthy trend and improve dietary choices. Nutrition andbiodiversity links directly to two of the Millennium DevelopmentGoals: to halve the proportion of people who suffer fromhunger; and to ensure environmental sustainability.7 However, thecontribution of biodiversity and underutilized species to nutritionand health has long been neglected. The nutrient potential ofmilk from a wide range of species and breeds has not beenfully explored. Acknowledging the importance of biodiversity forimproved nutrition is crucial for addressing malnutrition and for

appropriate nutrition interventions. Developing the biodiversityknowledge base also provides motivation to maintain local speciesand breeds which may otherwise become extinct; for example,in Italy, reductions in some autochthonous donkey populationssuch as the Ragusana breed and di Pantelleria breeds have beenreported,8 while mechanization of many agricultural practices hasled to the decline of the mare breed Murghes, now threatenedwith extinction.9

Buffalo milk is ranked second in the world in production,contributing towards ∼13% of the world’s milk production.4

In the Himalayan region, water buffalo (Bubalus bubalis) is theprimary source of income earned from livestock and yields 98%of the total milk consumed by the population (Singh, 1992, citedin Meena et al., 2011).10 In Pakistan and India, buffalo accountsfor 63% and 56%, respectively, of the total milk production.4 Inthe mountainous regions of China, Mongolia, Russia, Nepal, India,Bhutan, Tajikistan and Uzbekistan, where no other bovines arereared, the populations rely heavily on yak (Bos grunniens) formilk, meat, fur and transportation (Wiener, 2002, cited in Silk et al.,2006).11 Yak milk is dried in several factories in China, Nepal andMongolia for domestic consumption.12 Mithun (Bos frontalis), adomesticated bovine species, is mainly found in the hill regionsof India, Myanmar, Bhutan and Bangladesh,13 where it plays an

∗ Correspondence to: Ute Ruth Charrondiere, Nutrition and Consumer ProtectionDivision, Food and Agriculture Organization of the United Nations (FAO), Vialedelle Terme di Caracalla 00153 Rome, Italy. E-mail: [email protected]

NutritionandConsumerProtectionDivision,FoodandAgricultureOrganizationof the United Nations (FAO), 00153 Rome, Italy

J Sci Food Agric (2011) www.soci.org c© 2011 Society of Chemical Industry

www.soci.org E Medhammar et al.

important role in the economic, social and cultural life of thelocal people. This animal has the possibility of being further usedfor milk production in order to improve the rural economy andnutrition status in these countries. Mithun cross cattle hybridsare also used as milk animals in parts of northeastern India andBhutan.14

It is estimated that approximately 30 million people across theworld drink mare (Equus ferus caballus) milk.15 Herds are mainlyfound in Russia, Kazakhstan, Kyrgyzstan, Tajikistan, Uzbekistan,Mongolia, and eastern and central Europe. Both mare and donkeymilks are similar to human milk in total protein and lactosecontents, have similar fatty acid and protein profiles and a fairlylow mineral content. Therefore, equine milks could be a betteralternative to cow milk for infants, especially those with cow milkallergies.8,15 – 17

In arid and semi-arid areas where cows are affected by theheat and lack of water and feed, camelids play a major role insupplying the population with milk.18 There are two species ofcamel: the dromedary or Arabian camel (Camelus dromedaries)and the Bactrian camel (Camelus bactrianus). Out of an estimated18 million camels in the world, only 2 million are Bactrian camels.19

In Somalia, Mali and Ethiopia, camel milk represents about 40%,17% and 12%, respectively, of the total milk production.4 Llama(Lama glama) and alpaca (Lama pacos) are domesticated SouthAmerican camelids. Unlike camels, they have historically not beenbred for dairy purposes and information on milk composition andconsumption is scarce. Llama and alpaca milks are still under-exploited nutritional and economic resources for the millions ofpeople living in the mountainous areas of South America.17,20

Reindeer (Rangifer tarandus) herding is practised by about 20ethnic groups, from the Saami people of northern Scandinavia andthe Kola Peninsula, to the Koryak and Chukch people of easternSiberia. Reindeer milking is labour intensive and the task usuallyfalls to women.21 Renewed interest in reindeer milk lies in theexpanding market of gourmet products.21

Moose (Alces alces), also known as European elk, is the world’slargest antlered mammal and can be found in Canada, Alaska,Russia, Sweden, Norway, Finland and the Baltic states. Moosemilking farms, e.g. the Kostroma moose farm,22 can be found inRussia.

Minor dairy animals have considerable potential in contributingto food and nutrition security. They have been domesticated andbred for dairy purposes in regions where harsh environments andclimates require animals with special adaptations. They contributeto sustainable family nourishment, assure an income for therural population, enhance agricultural development, preserve localresources and improve human nutrition within these regions.23

Emphasizing the role of dairy animals other than cow, goatand sheep in milk production contributes directly to biodiversityenhancement by the preservation of traditional species, productsand local culture.

It is well known that milk composition varies among species,breeds within the same species, and even among individualanimals within the same breed.24,25 There are many reasons forthese differences in milk composition, including stage of lactation,breed differences, number of calvings (parity), seasonal variations,age and health of animal, feed and management effects includingnumber of milkings per day and herd size.24,26 – 28 The knowledgeof differences in nutrients and other bioactive components in milkamongst species and breeds allows the development of productsfor consumers with special needs, e.g. alternatives to cow milk for

people with allergy and milk intolerance,12,29 and improved milkcomposition through selection and cross-breeding.

The nutritional value of milk of minor dairy animals in humannutrition has so far received little attention in research and needs tobe emphasized. Previous reviews of milk from goat, sheep, cameland mare have presented different aspects in milk compositionamongst one or two species,30 – 32 but to our knowledge no reviewpaper on the milk composition at breed level of such a largenumber of species, which also includes data on underutilizedspecies, is available. The aim of this review is therefore to report,compare and highlight the inter- and intraspecies differences andsimilarities in composition of milk from various minor dairy animalsand buffalo.

EXPERIMENTAL (COMPILATION OF NUTRIENTCOMPOSITION DATA)Literature searchRelevant articles were obtained through an extensive literaturesearch in reference databases (using Scopus, Science Direct andCab Abstracts), book chapters, nutrition and agricultural journalsduring the period of November–January 2009/2010. Keywordsused were: milk, breed, biodiversity, milk AND the different animalspecies. Each bibliography of references generally led to severalother papers.

Only milks of species known to be consumed by humans wereincluded. Information on dairy products was not compiled. Dataon milk composition less then 30 days from parturition were nottaken into account, with the exception of one entry for moose milkfrom 15 to 25 days after parturition, and one entry for reindeermilk from lactation week 3. These were included because of thelimited data available for moose and reindeer milk and becausethe values were very similar to other values.

Information was gathered on the composition of whole rawmilk of dairy animals at the breed level, or at species level foranimals classified as underutilized. Breed is referred to as ‘asubspecific group of animal species, within a single zoologicaltaxon of the lowest known rank’.33 As the same breed may bereferred to by different names in different countries or regions,care was taken to identify such cases. Yak, Bactrian camel,donkey, moose, reindeer, musk ox, llama, alpaca and mithunwere categorized as underutilized species in regard to milkproduction, meaning that they are ‘species with underexploitedpotential for contributing to food security, health and nutrition’.33

They were identified taxonomically at the species level, andbreed name was noted when available. There were 119 scientificpapers that met the inclusion criteria, covering 271 data points.Of these, 13 papers were excluded because of problems withthe values or documentation. For three papers34 – 36 the valuesgiven far exceeded the normal range (e.g. one paper reportedα-tocopherol values in milk which were more than 1000 timeshigher than the average for the species); therefore these paperswere also excluded. Very large values were found for copper andmanganese,37 the value for the latter being more than 2000 timesthe value for cow (8147 µg compared with 4 µg 100 g−1) andmore than 45 times the highest value reported for any species(dromedary, 180 µg 100 g−1),38 and for this reason these valuestoo were excluded. The vitamin C values in one paper on camelmilk39 were excluded due to sample handling. For papers wherecontact information was provided, authors were contacted toobtain clarification prior to excluding a paper. Only one paperwith values for barren ground caribou was found,2 but had to be

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Composition of milk from minor dairy animals and buffalo breeds www.soci.org

excluded as the values were for milk obtained within 1 week ofparturition.

Buffalo have historically been divided into swamp and riverbuffalo based on morphological, behavioural and geographicalcriteria.40 The two types also differ in chromosome number, withswamp buffalo having 48 chromosomes and river buffalo 50chromosomes (Ulbrich and Fischer, 1967, cited in Groeneveldet al., 2010;40 Fischer and Ulbrich, 1968, cited in Groeneveld et al.,201040). They are sometimes referred to as different subspecies:river buffalo as Bubalus bubalis bubalis and swamp buffalo asBubalus bubalis carabenesis. Swamp buffalo have no recognizedbreeds.40 They are reported to be mainly used as draught animals41

and their milk yield is poor.10 Conversely, river buffalo are mainlyused for milk production.42 In the present work, we differentiatedby buffalo breeds and not by subspecies, and it is likely that ourdata are mainly limited to river buffalo breeds.

Camels may also be classified according to their function (racing,draught or dairy), or their habitat (lowland/mountain).19 However,in the present work we differentiated among camels only at specieslevel (dromedary versus Bactrian) and breed level.

We did not attempt to classify animals according to milk yieldsor maternal body weight in the present study. Although lactation-averaged values were used when they were presented in somepapers, other papers did not report lactation-averaged values butonly individual values for various lactation stages; in such cases,all reported values were used.

Standardization of dataAmong the data sources, considerable differences were encoun-tered in data expression. Protein and fat were sometimes reportedper 100 g milk, per 100 g protein or 100 g fat, or per 100 g aminoacids or per 100 g fatty acids. Units for micronutrient composi-tion were expressed in mg 100 g−1, mg kg−1, µg g−1, ppm andmmol L−1. Therefore, the compilation of macronutrients and mi-cronutrients required standardization prior to any comparisons.Differences in the data presented in this paper and the originalsource are a result of these recalculations. Although the majorityof data were from primary publications, sometimes it was notpossible to refer back to the original source and values cited in thepapers had to be used.

Data were recalculated to grams, milligrams or microgramsper 100 g fresh milk. When the concentration (mmol L−1) wasgiven, the values were recalculated to mg 100 g−1 by using thecorresponding molecular weight. Values for amino acids that weregiven per 100 g protein were recalculated to 100 g milk. Valuesfor fatty acids that were given per 100 g fat or fatty acids wererecalculated to 100 g milk, by using the conversion factor 0.945.43

In several papers presenting amino acids or fatty acids, no proteinor fat values were reported. Therefore average protein and fatvalues were calculated and applied per species, to express aminoacids and fatty acids per 100 g milk. When no nitrogen conversionfactor for protein was indicated, the factor 6.38, recommendedfor milk and milk products,44 was used. Internal checks on thenutrient profiles were carried out as recommended by Greenfieldand Southgate.43 Mean values were calculated when only a rangewas given in the scientific papers. Where breed-level informationwas available, average values per breed were used. Some of thesedata included different feeding regimes and lactation stages. Theonly exceptions were individual fatty acids, where data werenot averaged for breed, for reasons discussed under ‘‘Interbreeddifferences in milk composition’’.

Some papers reported values for folic acid, but since folate isthe naturally occurring form of the vitamin it was assumed to befolate. It was impossible to differentiate among all the active formsof vitamins A and E. Thus they were simply reported as vitaminA or vitamin E. Values for L-ascorbic acid were grouped togetherwith vitamin C.

The percentage of cow, goat, sheep, camel and buffalo milkproduction was obtained from FAO STAT 2008,4 using the valuesfor milk production per species and total milk production.

The amount of data available varied with species. A briefsummary of the data collected is given in Table 1.

Cow milkCow milk (the term ‘cow’ is used here to refer to the female ofBos taurus and Bos indicus) was included in this study as a genericreference for comparison purposes. Nutrient data for whole milk(3.25% milk fat, without added vitamin A and vitamin D, USDAcode 01211) were obtained from the USDA National NutrientDatabase45 and are included in Table 2 and in the nutrient tables.These values were chosen as a reference because the number ofdata points from which they had been derived were sufficientlylarge (ranging from 12 to 46 for the main nutrients).

Statistical analysisA statistical analysis using SPSS for MAC (version16.0) was carriedout on proximate data and mineral data. For the other nutrients, notenough data points were available after the data were averagedaccording to breed. The averaging was done in order to give anequal weight to data from all breeds, as otherwise the mean valuewould have been biased by data from the more commonly studiedbreeds, which contribute more data points. Data are presentedas mean ± SD. To account for significant differences among theanimal species for protein, total fat, ash, lactose, water, calcium,magnesium, phosphorous and potassium, the one-way ANOVA(followed by the Sidak) test was applied. Significance was set atP < 0.05.

RESULTS AND DISCUSSIONInterspecies differences in nutrient compositionTable 2 shows proximate data for all the species and includes theresults of the statistical analysis for buffalo, yak, mare, donkey,dromedary camel and reindeer milks; milk from the other speciesdid not have enough data points for inclusion in this analysis. Inthe following sections, the values referred to are for the averagecomposition of the milk, unless otherwise mentioned.

Comparison with cow milkBovine species buffalo (Bubalus bubalis), yak (Bos grunniens) andmithun (Bos frontalis). Milk from yak and mithun contained moreprotein on average than buffalo and cow milks. The highestreported protein value was 6.8 g 100 g−1 in mithun milk – data forlate lactation.46 Yak and buffalo milks were significantly differentfrom each other (P < 0.05) only in their total protein content.The milk from all three species contain appreciably higher valuesfor average total fat than cow milk, with the maximum individualvalue reported being 10.3 g 100 g−1 for mithun milk – data forlate lactation.46 This value is unusually high, considering thatthe mithun is not an Arctic animal, although its habitat is at anelevation of 1000–3000 m above sea level. The high values for fatand protein in mithun milk have been attributed to the low milkyield.46,47

J Sci Food Agric (2011) c© 2011 Society of Chemical Industry wileyonlinelibrary.com/jsfa


Tab

le1

.Su

mm

ary

ofd

ata

colle

cted

on

milk

com

po

siti

on

Spec

ies

Nu

mb

ero

fdat

ap

oin

tsN

um

ber

ofb

reed

sre

po

rted

Co

un

try

ofo

rig

ino

fdat

aN

utr

ien

tsfo

rwh

ich

no

dat

aw

ere

rep

ort

ed

Bu

ffal

o75

fort

ota

lfat

,up

to19

entr

ies

ind

ivid

ual

fatt

yac

ids,

1en

try

each

fori

nd

ivid

ual

amin

oac

ids,

vita

min

san

dso

me

min

eral

s.

18A

rgen

tin

a,B

razi

l,B

ulg

aria

,Ch

ina,

Fran

ce,G

erm

any,

Ind

ia,I

taly

,Ja

pan

,Nep

al,P

akis

tan

WP,

Se,M

n,v

itam

inD

Yak

54fo

rto

talf

at,1

dat

ap

oin

tfo

rso

me

ind

ivid

ual

fatt

yac

ids

and

amin

oac

ids

19M

on

go

lia,C

hin

a,In

dia

,Tib

et,N

epal

,Ru

ssia

,Afg

han

ista

n,K

yrg

yzst

anV

itam

ins,

Se,I

,Mn

,Cl,

WP

Mit

hu

n6

forc

asei

nan

dto

talp

rote

in,4

forw

ater

,to

tal

solid

s,to

talf

atan

dla

cto

se,2

forw

hey

pro

tein

and

1ea

chfo

rash

,so

lids

–n

ot

fat,

calc

ium

and

ph

osp

ho

rus

No

bre

eds

Ind

iaV

itam

ins

(all)

,AA

,FA

,Fe,

Mn

,K,N

a,Z

n,C

u,S

e,I,

Mn

,Cl

Mar

e44

fort

ota

lfat

,18

–26

form

ain

nu

trie

nts

30b

reed

s,w

hic

hin

clu

ded

the

Mo

ng

olia

nw

ildh

ors

eb

reed

Prze

wal

ski

Ital

y,Po

rtu

gal

,Hu

ng

ary,

Pola

nd

,Ru

ssia

,USA

AA

,Se,

Mn

,vit

amin

s(A

,E,D

,B5,

B6,

B12

,B9,

B7)

Do

nke

y12

fort

ota

lfat

,11

fort

ota

lpro

tein

and

lact

ose

,9fo

rwat

eran

dto

tals

olid

s,an

d3

forc

asei

n,w

hey

pro

tein

,lys

ozy

me,

β-l

acto

glo

bu

linan

dα

-lac

talb

um

in.1

–2

dat

ap

oin

tsfo

ram

ino

acid

s,M

g,K

,Na,

Cl,

and

vita

min

san

dre

po

rted

ener

gy,

calc

ium

,ph

osp

ho

rus,

lact

ofe

rrin

and

seru

mal

bu

min

.2–

3d

ata

po

ints

fori

nd

ivid

ual

fatt

yac

ids

4b

reed

s:Ra

gu

san

a,M

arti

na

Fran

ca,J

ian

gyu

ean

dLi

tto

ral-

Din

aric

Ital

y,C

hin

a,C

roat

ia,U

SAFe

,Zn

,Cu

,Se,

I,M

n,V

itam

ins

(A,E

,D

,C,B

5,B

6,B

12,B

7,B

9)

Dro

med

ary

cam

el14

dat

ap

oin

tsfo

rlac

tose

,7d

ata

po

ints

forv

itam

inC

and

4d

ata

po

ints

form

ajo

rnu

trie

nts

.1d

ata

po

int

each

form

ost

fatt

yac

ids

17Su

dan

,Ken

ya,T

un

isia

,In

dia

,K

azak

hst

an,C

hin

a,Sa

ud

iAra

bia

,Is

rael

,Ch

ile,P

eru

,Arg

enti

na,

USA

Vit

amin

s(e

xcep

tC

and

B2)

,AA

,WP,

Se,I

,Cl

Bac

tria

nca

mel

1–

2d

ata

po

ints

on

ly,f

orp

roxi

mat

es,f

atty

acid

s,m

ilkp

rote

ins,

afe

wvi

tam

ins

and

min

eral

s1

bre

ed(A

lxa)

Kaz

akh

stan

,Ch

ina/

Inn

erM

on

go

liaA

A,W

P,C

,Fe,

Cu

,Se,

I,M

n,v

itam

ins

(B3,

B5,

B12

,B9)

,B7

Alp

aca

2–

3fo

rto

talp

rote

in,t

ota

lfat

,lac

tose

,wat

eran

das

h1

bre

ed(H

uac

aya)

Ch

ile,P

eru

(On

lyp

roxi

mat

ed

ata

avai

lab

le)

Llam

a3

–4

forw

ater

,to

talp

rote

in,t

ota

lfat

,ash

and

lact

ose

,2fo

rrep

ort

eden

erg

yan

dca

lciu

m,1

each

fori

nd

ivid

ual

fatt

yac

ids

No

bre

eds

Ch

ile,P

eru

,Arg

enti

na,

USA

Vit

amin

s(a

ll),A

A,N

,Fe,

Zn

,Cu

,Se,

I,M

n,C

l

Rein

dee

r24

fort

ota

lfat

,16

forl

acto

se,1

–3

dat

afo

ram

ino

acid

s,fa

tty

acid

san

dm

iner

als

No

bre

eds

No

rway

,Sw

eden

,Fin

lan

d,A

lask

a.Ru

ssia

Vit

amin

s(a

ll),F

e,C

u,S

e,I,

Mn

,WP,

N

Mo

ose

8fo

rto

talp

rote

in,1

fors

om

eo

fth

efa

tty

acid

san

dto

taln

itro

gen

.7fo

rwat

er;6

forc

alci

um

,m

agn

esiu

m,s

od

ium

,po

tass

ium

;5fo

rto

talf

at,

lact

ose

and

ash

2b

reed

sre

cord

ed,

‘do

mes

tica

ted

Taig

am

oo

se’

and

Ala

skan

mo

ose

(Alc

esal

ces

giga

s)

USA

(Ala

ska)

,Ru

ssia

Vit

amin

s(a

ll),A

A,I

,WP,

C

AA

,am

ino

acid

s;C

,cas

ein

s;C

l,ch

lori

de;

Cu

,Co

pp

er;F

A,f

atty

acid

s;Fe

,iro

n;I

,io

din

e;K

,po

tass

ium

;Mn

,man

gan

ese;

N,N

itro

gen

;Na,

Sod

ium

;Se,

Sele

niu

m;Z

n,z

inc;

WP,

wh

eyp

rote

ins.



Tab

le2

.Pr

oxi

mat

eco

mp

osi

tio

no

fmilk

fro

mth

eva

rio

us

spec

ies

(ave

rag

ean

dra

ng

e)

Per1

00g

milk

Co

wa

Bu

ffal

ob

Yak

cM

ith

un

dM

usk

oxe

Mar

efD

on

keyg

Dro

med

ary

cam

elh

Bac

tria

nca

mel

iLl

amaj

Alp

acak

Rein

dee

rlM

oo

sem

Ener

gy,

calc

ula

ted

∗A

vera

ge

263

(63)

412

(99)

417

(100

)51

0(1

22)

356

(85)

199

(48)

156

(37)

234

(56)

319

(76)

326

(78)

299

(71)

819

(196

)53

8

valu

e,kJ

(kca

l)(1

29)

Ran

ge

296

–49

533

5–

557

458

–57

517

1–

295

135

–21

518

5–

332

258

–35

823

7–

351

525

–10

79

(71

–11

8)(8

0–

133)

(110

–13

8)(4

1–

71)

(32

–51

)(4

4–

79)

(62

–86

)(5

7–

84)

(126

–25

8)

Ener

gy,

Rep

ort

edva

lue,

kJ(k

cal)

Ave

rag

e25

6(6

1)36

8(8

9)19

3(4

6)21

0(5

0)39

2(9

4)88

0(2

09)

Ran

ge

349

–38

217

7–

210

388

–39

668

0–

1139

(87

–91

)(4

2–

50)

(93

–95

)(1

62–

272)

Wat

er(g

)A

vera

ge

88.1

83.2

a82

.6a

78.6

83.6

89.8

b90

.8b

89.0

b84

.884

.883

.767

.9c

76.8

Ran

ge

87.7

–89

.282

.3–

84.0

75.3

–84

.477

.4–

79.7

87.9

–91

.389

.2–

91.5

88.7

–89

.483

.7–

86.9

83.2

–84

.261

.9–

76.3

74.3

–79

.2

Tota

lpro

tein

(g)

Ave

rag

e3.

24.

0a5.

2b6.

55.

32.

0c1.

6c3.

1d3.

94.

15.

810

.4e

10.5

Ran

ge

3.1

–3.

32.

7–

4.6

4.2

–5.

96.

1–

6.8

1.4

–3.

21.

4–

1.8

2.4

–4.

23.

6–

4.3

3.4

–4.

33.

9–

6.9

7.5

–13

.07.

8–

14.4

Tota

lfat

(g)

Ave

rag

e3.

37.

4a6.

8a8.

95.

41.

6b,e

0.7b

3.2e

5.0

4.2

3.2

16.1

c8.

6

Ran

ge

5.3

–9.

05.

6–

9.5

7.7

–10

.30.

5–

4.2

0.3

–1.

82.

0–

6.0

4.3

–5.

72.

7–

4.7

2.6

–3.

810

.2–

21.5

7.0

–10

.0

Lact

ose

(g)

Ave

rag

e5.

14.

4a4.

8a4.

44.

16.

6b6.

4b4.

3a4.

26.

35.

12.

9c2.

6

Ran

ge

4.9

–5.

63.

2–

4.9

3.3

–6.

24.

1–

4.6

5.6

–7.

25.

9–

6.9

3.5

–4.

95.

9–

6.5

4.4

–5.

61.

2–

3.7

0.6

–3.

6

Ash

(g)

Ave

rag

e0.

70.

8a0.

8a0.

91.

60.

4b0.

4b0.

80.

90.

71.

61.

5c1.

6

Ran

ge

0.7

–0.

80.

4–

1.0

0.3

–0.

50.

3–

0.4

0.6

–0.

91.

4–

1.7

1.2

–2.

71.

5–

1.6

∗ Ap

pro

xim

ate

valu

eca

lcu

late

db

yu

sin

gth

een

erg

yva

lue

forp

rote

in(1

7kJ

),la

cto

se(1

6kJ

),an

dfa

t(3

7kJ

).Th

eva

lues

inkc

alw

ere

ob

tain

edb

yu

sin

gth

eco

nve

rsio

nfa

cto

rs1

kJ=

0.23

9kc

al.

Tab

le2

incl

ud

esth

ere

sult

so

fth

est

atis

tica

lan

alys

isfo

rb

uff

alo

,yak

,mar

e,d

on

key,

dro

med

ary

cam

elan

dre

ind

eer

milk

s;th

eo

ther

milk

sd

idn

ot

hav

een

ou

gh

dat

ap

oin

tsfo

rin

clu

sio

nin

this

anal

ysis

.V

alu

esin

aro

ww

ith

diff

eren

tle

tter

sar

esi

gn

ifica

ntl

yd

iffer

ent

(P<

0.05

).a

Co

w.U

SDA

.45

bB

uff

alo

.A

vera

ge

valu

esfr

om

Mu

rrah

,10,4

2,70

–72

,88,

95–

103,

129,

133,

134

Egyp

tian

bu

ffal

o,57

Zaf

arab

adi,95

Meh

san

a,95

,98

Bu

lgar

ian

Mu

rrah

,97,1

04,1

05B

ulg

aria

nb

uff

alo

×M

urr

ahb

reed

,97B

had

awar

i,98N

iliRa

vi,41

,42,

86Ja

ffra

bad

i,Su

rti,

Nag

pu

ri,P

and

har

pu

ri,a

nd

Tod

a,98

Ku

nd

i,41K

utt

anad

dw

arfb

uff

alo

,106

Zaf

farb

adia

nd

no

n-d

escr

ipt

hill

bu

ffal

o,10

Mu

rrah

×M

edit

erra

nea

n(c

ross

-bre

ed),96

,107

Med

iter

ran

ean

bu

ffal

o.10

8,10

9

cY

ak.A

vera

ge

valu

es,2,

11,1

10–

117

Him

ach

aliy

ak,11

8Pa

mir

,Kaz

aski

stan

iyak

,Mo

ng

olia

nya

kan

dU

pp

erA

ltai

yak,

119

Go

rno

-Alt

ai,57

Kir

gh

iz,57

,119

Zo

mo

(yak

×ze

bu

hyb

rid

s),11

8K

hai

nag

,113

Mai

wa,

37,1

17,1

19,1

20

Tian

zhu

Wh

ite,

117,

119,

120

Jiu

lon

g,11

5,11

7,12

0D

imzo

Jom

(cro

ss-b

reed

)an

dU

ran

gJo

m(c

ross

-bre

ed),11

Sib

u,43

Pali

and

Jial

i,117,

120

Gu

olu

o,H

uan

hu

,Zh

on

gd

ian

.117

dM

ith

un

.Ave

rag

eva

lues

.46,4

7

eM

usk

ox:

Ten

er(1

956)

cite

din

ref.

2.f

Mar

e.A

vera

ge

valu

esfr

om

Kir

gh

iz,

New

Kri

gh

iz,

Imp

rove

dK

irg

hiz

,tr

ott

ers,

Kir

gh

iztr

ott

er,

Kaz

akh

,B

ash

kir,

Russ

ian

bre

edo

fm

ilkin

gm

ares

and

San

am

ares

(mts

yri),

57M

urg

ese,

9,12

1It

alia

nsa

dd

leh

ors

e,12

2,13

5Lu

sita

no

,93,9

4H

aflin

ger

,77,8

2,92

,137

pri

mit

ive

Ko

nik

ho

rse,

85W

ielk

op

ols

ka,74

Perc

her

on

,123,

131

sad

dle

po

ny,

59B

ard

igia

no

,122

Palo

min

o,13

1Th

oro

ug

hb

red

,124,

138

Shet

lan

dp

on

y,12

4,13

8B

reto

n,

Bro

ulo

nn

ais

and

Hu

ng

aria

nD

rau

gh

t,82Q

uar

tera

nd

Rap

idH

eavy

Dra

ft,77

Prze

wal

ski.83

,124

gD

on

key.

Ave

rag

eva

lues

,2,56

,83,

124

Rag

usa

na,

8Li

tto

ral-

Din

aric

ass,

84Ji

ang

yue,

80M

arti

na

Fran

ca,13

6m

ixo

fmilk

fro

mRa

gu

san

aan

dM

arti

na

Fran

cab

reed

.81

hD

rom

edar

yca

mel

.Ave

rag

eva

lues

fro

mW

adh

a,39

,125

Afr

a,M

uja

hee

nB

lack

,Mal

ahB

lack

and

Safr

a,39

Ham

ra,39

,125

Maj

ahei

m,36

,63,

125

Som

ali,

Turk

ana,

and

Som

ali×

Turk

ana,

87Ja

isal

mer

i,B

ikan

eri,

and

Kac

hch

hi,12

6N

ajd

i,38,7

8A

rvan

a.32

iB

actr

ian

cam

el.A

vera

ge

valu

es,2

Alx

a.65

jLl

ama.

Ave

rag

eva

lues

.17,2

0,51

,91,

130

kA

lpac

a.A

vera

ge

valu

es,20

Hu

acay

a.91

lRe

ind

eer.

Ave

rag

eva

lues

,21,5

3,55

,57,

139

Ala

skan

rein

dee

r.139

mM

oo

se.A

vera

ge

valu

es,2,

23,5

2,12

8d

om

esti

cate

dTa

iga

mo

ose

.140



Musk ox (Ovibos moschatus). The musk ox is an Arctic mammalthat belongs to the subfamily Caprinae, as do goat and sheep.Only data on proximate composition were available for musk ox,obtained from one study (Tener, 1956, cited in Alston-Mills, 1995).2

Although the milk contains more fat than cow milk, the fat content(5.4 milk g 100 g−1) is not high for an Arctic animal.

Equine species mare (Equus ferus caballus) and donkey (Equusafricanus asinus). The milk from these two non-ruminant specieswas very similar to each other: no significant differences (P < 0.05)were seen in protein, fat, lactose, ash or water contents. Both maremilk and donkey milks contained substantially lower amounts oftotal fat and total protein compared with cow milk. The lactose andwater contents of both mare and donkey milks were higher thanof cow milk. The values for ash were nearly half the correspondingvalue for cow milk. Equine milks have been reported to be nearestin composition to human milk because of the high lactose and lowprotein and mineral contents48 and fatty acid composition.49

Camelid species dromedary camel (Camelus dromedaries), Bactriancamel (Camelus bactrianus), llama (Lama glama) and alpaca (Lamapacos). Camelids have a stomach with three compartmentsrather than four but with similar functional properties toruminant stomachs;50 therefore they are sometimes called‘pseudoruminants’.

The lactose and protein contents in the milk from the two camelspecies were similar but their fat contents differed. Bactrian camelmilk varied from dromedary camel milk mainly in fat content: theaverage fat content of Bactrian camel milk was 5 g 100 g−1 milk. Inoverall proximate composition, dromedary camel milk was mostsimilar to cow milk. Alpaca milk was richer in protein and ashthan other camelid milks, having an average protein value of 5.8 g100 g−1 and an average value for ash of 1.6 g 100 g−1 – nearlydouble the values reported in the other milks. Llama milk had thehighest average lactose content in this group, with a maximumreported value of 6.3 g 100 g−1 milk.51

Cervid species reindeer (Rangifer tarandus) and moose (Alces alces).Both these species are noted for their concentrated milk, which hasa cream-like consistency.52,53 Both milks contained on average 7 g100 g−1 or more protein than cow milk. The average fat contentof reindeer milk was almost double that of moose milk and nearlyfive times the value for cow milk. Both reindeer and moose milkwere low in lactose, containing approximately half the value foundin cow milk.

Variations in proximate composition among speciesThe interspecies range between the lowest and highest averageprotein content was nearly 9 g 100 g−1, with the highest valuesreported in cervid milks and the lowest in equine milks. Just onecup (250 mL) of moose or reindeer milk (which provide on average26 g protein) equals the recommended safe level of protein intake(∼26 g per day) for children below 10 years of age.54

Reindeer milk, reported to be second only to marine mammalsin fat content,48 had by far the highest amount of average fat(16 g 100 g−1): the highest individual value reported for reindeermilk was 21.5 g 100 g−1.55 Donkey milk contained the least fat(0.7 g 100 g−1), while mare milk also had a very small average fatcontent (1.6 g 100 g−1), with the lowest value being 0.3 g 100 g−1

for donkey milk.56

The average lactose content was lowest in moose milk (2.6 g100 g−1) and reindeer milk (2.9 g 100 g−1). The Saami people, whoare reindeer herders, are reported to be rather intolerant of lactose;hence reindeer milk is particularly suitable.53

The highest average lactose content was in mare milk (6.6 g100 g−1); donkey and llama milks also had a high content oflactose, with average values of 6.4 and 6.3 g 100 g−1, respectively.The highest reported value for lactose was 7.2 g 100 g−1 in maremilk.57

Reported energy values were available for five of the speciesincluded in this study, and Table 2 includes both reported andcalculated values for energy. The differences between the reportedand calculated energy values can be attributed to a number ofcauses, including the fact that the calculated values are based onthe average values for protein, lactose and fat, and the reportedvalues may not have always included an analysis of all energy-yielding components. Some of the reported values may havebeen for total combustible energy, while others would have beencalculated energy values. The multiplicity of analytical methodsavailable for analysing protein, fat and carbohydrates and thevariety of different available food energy conversion factors canalso lead to different energy values.58

Reindeer milk provided the most energy – more than four timesthe amount from mare and donkey milks. In reindeer milk, the highenergy and protein may be optimal to meet both the growth andenergy requirements of the calf, which enables the calf to survivethe harsh Arctic winter; the concentrated milk is reported to beparticularly suitable for the locomotive lifestyle of the reindeer.55

Mithun and moose milks also provided more than 500 kJ 100 g−1

of energy.The average ash content was highest in musk ox, alpaca and

moose milks (1.6 g 100 g−1), while reindeer milk had a similarlyhigh value of 1.5 g 100 g−1. The highest individual value reportedwas for reindeer milk: 2.7 g 100 g−1. The lowest values were inmare and donkey milks: 0.4 g 100 g−1. In general, an inverserelationship between ash content and lactose content was seen,with milk containing the lowest ash content (mare and donkeymilks) containing the highest lactose content, and vice versa.This is as expected, as lactose, together with sodium, potassiumand chloride ions, plays a major role in maintaining the osmoticpressure in the mammary system.59

Within a species, differences in water content were not verylarge – generally 2–3% – with the exception of reindeer (14%) andyak (9%) milks. In order to see if the large variation in total proteincontent in reindeer milk merely reflected differences in watercontent, the total protein was calculated per 100 g dry weight(DW) basis for the lowest and highest protein values, using thecorresponding water values reported in those studies. The rangethus obtained was 31.6–34.1 g 100 g−1 DW, which still shows adifference of 2.5 g between the lowest and highest values. Whenthe values for total fat in yak milk were reviewed, the study givingthe lowest value (5.6 g 100 g−1) and the study giving the secondhighest value (8.6 g 100 g−1) had the same water content (84.4 g100 g−1), further supporting the conclusion that the range ofvalues does not just correspond to changes in water content.

MineralsThe amount of available data for minerals varied, depending onspecies (Table 3). Buffalo, moose, mare and dromedary camel datawere fairly complete, while data for yak, donkey, Bactrian camel,llama and reindeer were limited. Mithun data were found only for



Figure 1. Interspecies differences in mineral content.

Figure 2. Vitamin and mineral content for milk from various species in relation to the recommended nutrient intake values (RNI).

calcium and phosphorus. Sulfur data were found only for Bactriancamel (35.9 mg 100 g−1).60

Interspecies differences were seen for most elements (Fig. 1).Moose and reindeer milks contained the highest quantity of mostminerals, while mare and donkey milks contained the lowestquantities, which is a reflection of their respective ash contents.However, relative concentrations of minerals differed in the milkfrom the various species. For example, the calcium : phosphorusratio in moose milk and buffalo milk was almost 1 : 1; inmithun milk, this ratio was 1 : 0.6, while the calcium contentof milk from all the other species was 1.2–1.6 times morethan that of phosphorus. In general, a high molar ratio ofcalcium : phosphorus is recommended61 in order for the bodyto maintain adequate calcium levels and prevent bone resorption.The potassium : sodium ratio also varied in the milk from differentspecies. Most milks contained two to three times more potassium

than sodium, while llama milk contained four times morepotassium than sodium.

Figure 2 shows the recommended nutritional intake (RNI) ofminerals for a 1- to 3-year-old child62 versus the intake fromvarious milks. Two cups (500 mL) of milk from most species caneasily supply more than 70% of the RNI of calcium. Two cups aday of either reindeer or yak milks would supply the RNI of zinc,while Bactrian and dromedary camel milks and moose milk wouldprovide 70–90% of the RNI. The RNI for magnesium is provided bytwo cups of buffalo, dromedary camel, llama, reindeer and moosemilks, with two cups of moose milk providing 192% of the RNI.In contrast, two cups of donkey milk would provide only 33% ofthe RNI for this mineral, which clearly demonstrates the variationin magnesium content in milk from different species. None of themilks are a source of Fe. Even yak milk, which is the richest in iron,provides less than 60% of the RNI.



Tab

le3

.M

iner

alco

mp

osi

tio

no

fmilk

fro

mth

eva

rio

us

spec

ies

(ave

rag

e,w

ith

ran

ge

inp

aren

thes

es)

Per1

00g

milk

Co

wa

Bu

ffal

ob

Yak

cM

ith

un

dM

aree

Do

nke

yfD

rom

edar

yca

mel

gB

actr

ian

cam

elh

Llam

aiRe

ind

eerj

Mo

ose

kRN

Ilp

erd

ayfo

rch

ildre

n,

1–

3ye

ars

Cal

ciu

m(m

g)

113

191b

129a

,b88

95a

9111

4a,b

153.

719

532

028

0c50

0

(112

–11

6)(1

47–

220)

(119

–13

4)(7

6–

124)

(68

–11

5)(1

05–

120)

(152

.3–

155)

(170

–22

0)(1

56–

358)

Iro

n(m

g)

0.03

0.17

0.57

0.10

0.21

0.31

5(1

2%b

ioav

aila

bili

ty)

(0.0

0–

0.07

)(0

.15

–0.

98)

(0.0

3–

0.15

)(0

.17

–0.

26)

Mag

nes

ium

(mg

)10

12a

107a

413

a,b

8.1

1519

23b

60

(7–

12)

(2–

16)

(8–

12)

(4–

12)

(12

–14

)(1

9–

26)

Pho

sph

oru

s(m

g)

8418

5a10

6a,b

147

58b

6186

a,b

131.

512

227

027

6

(59

–95

)(1

02–

293)

(77

–13

5)(4

3–

83)

(49

–73

)(8

3–

90)

(117

–14

5.9)

Pota

ssiu

m(m

g)

132

112

9551

a50

151b

186.

112

015

611

1a,b

(106

–16

3)(8

3–

107)

(25

–87

)(1

24–

173)

(181

.2–

191)

(82

–15

0)

Sod

ium

(mg

)43

4729

1622

6666

.427

.248

78

(26

–58

)(2

1–

38)

(13

–20

)(5

9–

73)

(60.

8–

72)

(46

–50

)(3

7–

158)

Zin

c(m

g)

0.4

0.5

0.9

0.2

0.6

0.7

1.1

0.6

4.1

(mo

der

ate

bio

avai

lab

ility

)

(0.3

–0.

5)(0

.7–

1.1)

(0.2

–0.

3)(0

.4–

0.6)

Co

pp

er(m

g)

0.03

0.02

0.41

0.05

0.15

0.29

(0.0

0–

0.08

)(0

.02

–0.

11)

(0.1

2–

0.17

)

Sele

niu

m(µ

g)

3.7

1117

(0.0

–8.

8)

Iod

ine

(µg

)4

75

Man

gan

ese

(µg

)4

106

1

(0–

8)(6

0–

180)

Ch

lori

de

(mg

)57

1934

152

6880

Tab

le3

incl

ud

esth

ere

sult

so

fth

est

atis

tica

lan

alys

isfo

rbu

ffal

o,y

ak,m

are,

dro

med

ary

cam

elan

dm

oo

sem

ilks;

the

oth

erm

ilks

did

no

thav

een

ou

gh

dat

ap

oin

tsfo

rin

clu

sio

nin

this

anal

ysis

.Val

ues

ina

row

wit

hd

iffer

ent

sup

ersc

rip

tsar

esi

gn

ifica

ntl

yd

iffer

ent

(P<

0.05

).a

Co

w.U

SDA

.45

bB

uff

alo

.Ave

rag

eva

lues

fro

mEg

ypti

anb

uff

alo

,57K

utt

anad

dw

arfb

uff

alo

,106

Meh

san

aan

dZ

afar

abad

i,95M

urr

ah.95

,129

,134

cY

ak.A

vera

ge

valu

esfr

om

,111,

113

Kh

ain

ag(y

akh

ybri

d),11

3M

aiw

a.37

,115

Co

bal

t.111

dM

ith

un

.Ave

rag

eva

lues

.47

eM

are.

Ave

rag

eva

lues

fro

mIt

alia

nsa

dd

leh

ors

e,12

2,13

5H

aflin

ger

,137

Perc

her

on

,15B

ard

igia

no

,122

Palo

min

o,13

1Th

oro

ug

hb

red

,124,

138

Shet

lan

dp

on

y,12

4,13

8Pr

zew

alsk

iho

rse,

83,1

24Lu

sita

no

.94

fD

on

key.

Ave

rag

eva

lues

fro

m,83

mix

ofR

agu

san

aan

dM

arti

na

Fran

cam

ilk.81

gD

rom

edar

yca

mel

.Ave

rag

eva

lues

fro

mM

ajah

eem

,Wad

ah,H

amra

,125

Naj

di,38

Arv

ana.

64

hB

actr

ian

cam

el.A

vera

ge

valu

esfr

om

Alx

a.60

,65

iLl

ama.

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rag

eva

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Tab

le4

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itam

inco

nte

nt

ofm

ilkfr

om

the

vari

ou

ssp

ecie

s(a

vera

ge,

wit

hra

ng

ein

par

enth

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)

Per1

00g

milk

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wa

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ffal

ob

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ecD

on

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med

ary

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ele

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tria

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–3

year

s)

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amin

A(µ

g)R

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ge

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ean

req

uir

emen

t:40

0µg

RE/d

ay

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amin

E(m

g)(

α-t

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ph

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l)A

vera

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0.19

0.15

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vin

(mg

)A

vera

ge

0.17

0.11

0.02

0.03

0.06

0.12

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mg

/day

(0.1

6–

0.20

)(0

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–0.

03)

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cin

(mg

)A

vera

ge

0.09

0.17

0.07

0.09

6∗m

g/d

ay

(0.0

7–

0.12

)

Pan

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enic

acid

(mg

)A

vera

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0.15

2.0

mg

/day

(0.3

6–

0.38

)

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amin

B6

(mg

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vera

ge

0.04

0.33

0.05

0.5

mg

/day

(0.0

3–

0.04

)

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te(µ

g)

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160

µg/d

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(µg

)A

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0µg

/day

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)A

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0.40

0.9

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0.53

)

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C(m

g)

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ay

(1.7

–8.

1)(2

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18.4

)

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amin

D(µ

g)

1.6

5µg

/day

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ren

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um

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dm

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∗ mg

NE,

nia

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;RE,

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leq

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ts.



The low Fe content is one of the reasons why animal milks arenot recommended in the complementary feeding of infants below12 months.

VitaminsData for vitamins were limited, or absent altogether for severalof the species studied here. Available data are shown in Table 4.Dromedary camel milk had the highest average value for vitamin C(6.7 mg 100 g−1 milk), with the values ranging from 2.5 mg 100 g−1

(Majaheem breed)63 to 18.4 mg 100 g−1 milk (Arvana breed).64 Theavailability of even a moderate amount of vitamin C in camel milkis reported to have significant relevance to nutrition in areas wheregreen vegetable and fruits are hard to find (Sawaya et al., 1984,cited in Zhang et al., 2005).65 However, vitamin C in camel milk isreported to be more heat sensitive than in cow milk, decreasingby about 27% when the milk is pasteurized.63 While a value forvitamin C in cow milk is not available in the USDA database (0 mg100 g−1 milk), other databases (McCanes and Widdowson, DanishFood Composition Database and Colombian Food CompositionTable) report values from 1.2 to 2 mg 100 g−1 milk. Two cups ofdromedary camel milk can supply the RNI of vitamin C, while twocups of mare milk can supply 72% RNI (Fig. 2).

Buffalo milk had a vitamin B6 content of 0.33 mg 100 g−1, nearlyan order of magnitude larger than that of cow milk. Two cups ofbuffalo milk can provide 330% of the RNI (0.5 mg per day) of vitaminB6 for a 1- to 3-year-old child, while the same quantity of Bactriancamel milk and cow milk provide 50%RNI and 40% RNI, respec-tively. The RNI of riboflavin, 0.5 mg per day, can be provided by twocups of cow, buffalo or Bactrian camel milks, while mare, donkeyand dromedary milks can provide 20–60% of the RNI. Buffalo milkwas rich in biotin; even 100 g of milk can easily provide the RNI.Vitamin D is reported in Bactrian camel milk at 1.6 µg 100 g−1,65

with two cups of milk providing 160% of the RNI. However, forniacin and folate, two cups of milk a day from any of the dairyspecies listed in Fig. 2 are not able to provide even 20% of the RNI.

Amino acidsThe high-quality protein in milk plays a crucial role in nutrition,especially in the developing world where diets are largely cerealbased. Lysine is commonly limiting in wheat- or maize-based diets,being present at 57% and 58% of requirement levels.54 In cassava-based diets the branched-chain amino acids leucine, valine andisoleucine are the limiting amino acids, being present at 79% ofrequirement levels.54

Table 5(a) shows the distribution of amino acids as grams ofamino acid per100 g total protein. Table 5(b) shows the adultindispensable amino acid requirement,54 together with the aminoacid content of the milks expressed as a percentage of therequirement pattern values. It is clear that all milks are a richsource of indispensable amino acids, regardless of species.

It is useful to consider how the amino acid content of milk fromdifferent species compare, per equal volume of milk, as shown inTable 6. As expected, the amino acid content reflects the proteincontent; donkey milk contained the lowest amounts of AA andreindeer milk contained the highest, per 100 g milk. The lysinecontent of buffalo milk was double that of cow milk. However,the lysine value of standardized cow milk, 137 mg 100 g−1 milk,is lower than is usual for cow milk. For example, other values forcow milk in the USDA database vary from 246 to 287 mg 100 g−1

milk, while the value in the Danish Food Composition Database is310 mg 100 g−1 milk. Only yak milk (375 mg 100 g−1) and reindeermilk (909 mg 100 g−1) exceed these values.

Milk proteinsThe major milk proteins are caseins (αS1-, αS2-, β-, κ- andγ -caseins) and whey proteins, which include α-lactalbumin, β-lactoglobulin, serum albumin, lactoferrin, immunoglobulins andlysozyme. Although the content of milk proteins is generallyproportional to the total protein content, the pattern of milkproteins varies among species.

The casein fraction in total protein varies with species (Table 7).In mare and donkey milks it was on average 43% and 45% oftotal protein, respectively, while in dromedary and reindeer milkit was 86% and 80% of total protein, respectively. Reindeer milk,which has the highest protein content from the above species,contains nearly12 times as much casein per 100 g milk as donkeymilk, which contained the lowest average protein. The two equinemilks had the lowest amount of total whey proteins per 100 g milk,even though whey proteins make up 40–50% of the total proteinin these milks, compared with less than 20% in cow milk.56

Dromedary camel, Bactrian camel and llama milks are reportednot to contain measurable amounts of β-lactoglobulin;17,66,67 oneof the main proteins associated with cow milk allergy.

Fatty acidsTable 8 shows the fatty acid (FA) content per 100 g milk; for totalsubclasses of FA both reported values (given in the final rows of thetable) and calculated values, obtained by summing the reportedindividual FA, are given. Not all the individual FA were reportedin the papers; in some studies, only a limited number of the mainFA were analysed, while other papers only reported values for FAsubclasses, without giving values for individual FA. Table 9 showsthe FA profiles, expressed as g 100 g−1 total FA, for milk fromdifferent species, and includes some fatty acid subclasses andnutritionally important fatty acids.

Bovine (buffalo and yak). The FA profile in milk from buffalo andyak was similar to that of cow milk. The major FA in milk fromboth species were C14 : 0, C16 : 0, C18 : 0 and C18 : 1, as in cowmilk. Individual saturated fatty acids (SFA) have different effectson the concentration of lipoprotein cholesterol fractions, withC12 : 0, C14 : 0 and C16 : 0 increasing low-density lipoprotein (LDL)cholesterol, while C18 : 0 has no effect.68 Buffalo milk containedthe highest calculated amount of trans FA (average value 528 mg100 g−1 milk). Two cups a day of buffalo milk (500 mL) contains1.96–3.54 g trans FA. While trans FA from commercial partiallyhydrogenated vegetable oils can increase coronary heart disease(CHD) risk factors and CHD events, the estimated average ruminanttrans FA intake in most societies is low:68 the estimated averagedaily ruminant trans FA intake of American adults is 1.2 g,compared with an average total daily trans FA intake of 5.8 g.69

Milk from both buffalo and yak contained a small amount ofconjugated linoleic acid (CLA), (1 g 100 g−1 total FA for buffalo,0.2 g 100 g−1 total FA for yak), the content of which can beenhanced by dietary manipulations, e.g. inclusion of plant oils infeeds.70 – 72 The biologically active form of CLA is thought to beC18 : 2 cis-9,trans-11,73 which occurs almost exclusively in ruminantmilk fat.

Equine (mare and donkey). Most individual fatty acid values weresmallest in donkey and mare milks. Compared with cow milk,equine milks fat had a high content of polyunsaturated fatty acids(PUFA) (20 g 100 g−1 total FA of mare milk and 23 g 100 g−1 totalFA of donkey milk, compared with 6.3 g 100 g−1 total FA of cow



Table 5(a). Distribution of amino acids in milks as grams amino acid per 100 g total protein

Cowa Buffalob Yakc Donkeyd Reindeere

Average total protein (g 100 g−1 milk): 3.2 4.0 5.2 1.6 10.4

Amino acids: (g AA 100 g−1 protein)

Isoleucine 5.03 4.85 4.38 5.44 4.42

Leucine 8.13 9.20 12.23 8.44 9.34

Lysine 4.28 7.48 7.21 7.19 8.74

Methionine 2.28 2.33 2.75 1.75 2.61

Phenylalanine 4.50 4.58 4.48 4.25 4.48

Threonine 4.38 4.35 4.12 3.50 4.53

Tryptophan 2.28 1.63 1.38

Valine 5.88 5.85 6.02 6.38 5.83

Cysteine 0.50 0.83 0.98 0.44 0.75

Tyrosine 4.63 4.53 3.31 3.63 5.41

Arginine 2.28 2.55 3.79 4.50 2.97

Histidine 2.28 2.73 2.25 2.81

Alanine 3.16 3.03 2.63 3.44 3.08

Aspartic acid 7.25 7.13 6.69 8.75 6.42

Glutamic acid 19.8 21.4 16.6 22.4 20.3

Glycine 2.28 1.93 2.65 1.19 2.24

Proline 10.4 12.0 2.46 8.63 9.26

Serine 3.25 4.65 3.88 6.13 5.46

Total 92.6 99.4 85.8 98.3 100.1

a Cow.45

b Buffalo. Average values from Bulgarian Murrah.142

c Yak. Average values,11 from Khainag (yak hybrid,113 Maiwa.37

d Donkey. Average values from Jiangyue.80

e Reindeer. Average values,53 (Malinen et al., 2002; Holand et al., unpublished data; Luick et al., 1974).

Table 5(b). Distribution of amino acids in milks, expressed as a percentage of requirement pattern

Percentage of requirement patternRequirement pattern (adult indispensable

AA requirement∗) in mg g−1 protein Cowa Buffalob Yakc Donkeyd Reindeere

Isoleucine 30 168 162 146 181 147

Leucine 59 138 156 207 143 158

Lysine 45 95 166 160 160 194

Threonine 23 190 189 179 152 197

Tryptophan 6 380 272 231

Valine 39 151 150 154 163 149

Histidine 15 152 182 150 187

Phe + Tyr 38 240 239 205 207 260

Met + Cys 22 126 143 170 99 153

∗ ‘Protein and amino acid requirements in human nutrition’. Report of a joint WHO/FAO/UNU Expert Consultation. WHO technical Report series, p.935. Mean nitrogen requirement of 105 mg nitrogen kg−1 per day (0.66 g protein kg−1 per day).Met + Cys, methionine and cysteine, sulfur AA; Phe + Tyr, phenylalanine + tyrosine.a Cow.45


c Yak. Average values,11 from Khainag (yak hybrid),113 Maiwa.37



milk) and a low content of SFA, particularly mare milk, whichhad an average value of 38 g 100 g−1 total FA, compared with60 g 100 g−1 total FA in cow milk. There is convincing evidencethat replacing SFA (C12 : 0–C16 : 0) with PUFA decreases LDLcholesterol concentration and the total/high-density lipoprotein(HDL) cholesterol ratio.68

In addition, the milks contained α-linolenic acid (ALA) andlinoleic acid (LA), which are essential since they cannot besynthesized by humans.68 The highest values reported in mare milkwere 20 g 100 g−1 total FA for ALA in the milk from the primitiveKonik horse – an average for lactation months 1–674 – and 16 g100 g−1 total FA for LA in Wielkopolska mares – an average for



Table 6. Average amino acid composition (mg 100 g−1 milk) of cow, buffalo, yak, donkey and reindeer milks, with ranges in parenthesis

mg 100 g−1 milk Average Cowa Buffalob Yakc Donkeyd Reindeere

Isoleucine 161 194 228 87 460

(446–468)

Leucine 260 368 636 135 971

(958–987)

Lysine 137 299 375 115 909

(284–466) (804–1099)

Methionine 73 93 143 28 271

(243–295)

Phenylalanine 144 183 233 68 466

(211–254) (458–471)

Threonine 140 174 214 56 471

(206–223) (454–482)

Tryptophan 73 85 144

(132–155)

Valine 188 234 313 102 606

(287–339) (580–630)

Cysteine 16 33 51 7 78

(74–81)

Tyrosine 148 181 172 58 563

(550–573)

Arginine 73 102 197 72 309

(265–383)

Histidine 73 109 36 292

(255–363)

Alanine 101 121 137 55 320

(312–332)

Aspartic acid 232 285 348 140 668

(641–682)

Glutamic acid 634 857 864 358 2116

(1931–2319)

Glycine 73 77 138 19 233

(227–238)

Proline 334 481 128 138 963

(937–1011)

Serine 104 186 202 98 568

(554–580)

a Cow.45


c Yak. Average values,11 from Khainag (yak hybrid),113 Maiwa.37



lactation months 1–6,74 compared with 3.23 g 100 g−1 total FAfor LA in cow milk.75

The fatty acid composition in mare milk is reported to be similarto that of human milk,49 and particularly relevant for humannutrition.76 The high LA and ALA contents and low C18 : 0 areattributed to monogastric animals: the microbial hydrogenationof fatty acid in the digestive tract does not occur before intestinalabsorption in equines, as in the ruminant tract (Jenkins et al.,1996, cited in Chiofalo et al., 2001).8 Therefore, the high contentof unsaturated long-chain FA in equine milk is related to amountsconsumed with forage.8,77

It is interesting to note that although mare milk had the secondlowest content of total fat per 100 g milk, it contains the highestamount of total n-3 fatty acids per 100 g milk, an average of99 mg 100 g−1 and range of 58–307 mg 100 g−1, because of

the high content of ALA (average 94 mg 100 g−1). It has beensuggested that mare milk may be used in infant nutrition, as thelivers of infants are probably capable of transforming these FAinto eicosapentaenoic acid (EPA), docosahexaenoic acid (DHA)and arachidonic acid.76 The n-3 fatty acids EPA and DHA, as partof a healthy diet, can contribute to the prevention of coronaryheart disease (CHD) and possibly other degenerative diseasesassociated with ageing.68 For populations with no access to n-3 FAfrom fish, e.g. Mongolia, intake of mare milk is crucial for meetingrequirements (adequate intake of 100–150 mg EPA+DHA for a 2-to 4-year-old child; acceptable macronutrient distribution rangeof 0.25–2 g per day for adults).68

Camelids (dromedary camel, Bactrian camel and llama). Indromedary camel milk, the monounsaturated fatty acid (MUFA)



Tab

le7

.M

ilkp

rote

ins:

aver

age,

wit

hra

ng

ein

par

enth

eses

Per1

00g

milk

Bu

ffal

oa

Yak

bM

ith

un

cM

ared

Do

nke

yeD

rom

edar

yca

mel

fB

actr

ian

cam

elg

Llam

ahRe

ind

eeri

Mo

ose

j

Tota

lpro

tein

(g)

4.0

5.2

6.5

2.0

1.6

3.1

3.9

4.1

10.4

10.5

(2.7

–4.

6)(4

.2–

5.9)

(6.1

–6.

8)(1

.4–

3.2)

(1.4

–1.

8)(2

.4–

4.2)

(3.6

–4.

3)(3

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4.3)

(7.5

–13

.0)

(7.8

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.4)

Nit

rog

en,t

ota

l(g

)1.

00.

280.

290.

40.

81.

2

(0.2

6–

0.32

)(0

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0.5)

(0.5

6–

1.16

)

Wh

eyp

rote

in(m

g)

1505

790

760

1012

(142

0–

1590

)(6

65–

955)

(680

–85

0)

Cas

ein

(mg

)31

1030

6044

5085

571

426

5232

0083

00

(302

0–

3200

)(2

111

–38

00)

(427

0–

4770

)(8

35–

890)

(612

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0)(1

900

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90)

(800

0–

8600

)

β-C

asei

n(m

g)

1100

α-C

asei

n(m

g)

1200

Lyso

zym

e(m

g)

157

(100

–22

7)

Lact

ofe

rrin

(mg

)67

3021

22

(29

–31

)

β-L

acto

glo

bu

lin(m

g)

708

305

(534

–88

2)(2

03–

375)

α-L

acta

lbu

min

(mg

)19

817

734

0

(193

–20

4)(1

53–

197)

Imm

un

og

lob

ulin

(mg

)19

2383

64

Seru

mal

bu

min

(mg

)10

735

(79

–13

4)(2

8–

42)

aB

uff

alo

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rag

eva

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fro

mM

urr

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3an

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bY

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,143

Go

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37an

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ng

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cM

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7,14

4

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pid

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rag

eva

lues

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6,83

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e,80

mix

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aan

dM

arti

na

Fran

cam

ilk.81

fD

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mel

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eva

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adah

Ham

ra,a

nd

Maj

ahei

m,12

5an

dA

rvan

a.20

gB

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cam

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ge

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Alx

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7

hLl

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30

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ind

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eva

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39

jM

oo

se.A

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valu

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om

do

mes

tica

ted

Taig

am

oo

se.14

0



Tab

le8

.Fa

tty

acid

s(m

g10

0g

−1m

ilk)i

nm

ilkfr

om

the

vari

ou

ssp

ecie

s(a

vera

ge,

wit

hra

ng

ein

par

enth

eses

)

Co

wa

Bu

ffal

ob

n=

Bu

ffal

ob

Yak

c

(n=

1,ra

ng

eg

iven

wh

ere

n=

2)M

ared

n=

Mar

edD

on

keye

n=

Do

nke

yeD

rom

edar

yf

n=

Dro

med

aryf

cam

el

Bac

tria

ng

cam

el(n

=3

anim

als)

Llam

ahRe

ind

eeri

n=

Rein

dee

ri

Mo

ose

j

(n=

1,ra

ng

eg

iven

wh

ere

n=

2)

Tota

lfat

3.3

7.4

6.8

1.6

0.72

3.0

5.0

4.2

16.1

8.6

(g10

0g

−1

milk

)(5

.3–

9.0)

(5.6

–9.

5)(0

.5–

4.2)

(0.3

–1.

8)(2

.0–

6.0)

(4.3

–5.

7)(2

.7–

4.7)

(10.

2–

21.5

)

Fatt

yac

id(m

g1

00

g−1

milk

)

Satu

rate

dFA

C4

:075

829

49

12

31

386

395

336

(237

–38

0)(1

–2)

(2–

4)(5

10–

133)

(33

–38

)

C6

:075

1116

19

32

61

710

93

320

4

(114

–27

8)(1

–3)

(4–

7)(1

12–

474)

C7

:01

18

C8

:075

1111

513

322

612

596

381

696

(71

–18

5)(1

3–

49)

(46

–77

)(3

–7)

(28

–12

4)

C9

:01

352

C10

:075

1431

859

1365

289

38

114

410

923

1

(113

–12

24)

(31

–10

5)(6

7–

112)

(4–

13)

(61

–15

5)

C11

:01

12

C12

:077

1420

989

1373

238

329

2168

415

237

(75

–10

0)(1

47–

333)

(43

–11

5)(3

8–

38)

(25

–33

)(1

5–

24)

(93

–21

7)(2

4–

50)

C13

:07

22

1932

114

(1–

3)(1

4–

24)

C14

:029

714

922

419

1378

228

432

929

347

84

1720

210

(194

–33

4)(6

83–

1254

)(5

6–

110)

(21

–35

)(1

32–

476)

(233

–34

0)(1

190

–24

90)

(166

–25

3)

Cis

o14

:03

Tr1

11

C15

:03

168

667

72

24

4159

683

137

60

(155

–17

6)(5

–12

)(1

–2)

(17

–56

)(4

8–

74)

(103

–15

5)(5

0–

70)

Cis

o15

:04

2(1

–3)

229

(18

–41

)

Can

teis

o15

:02

61(5

1–

71)

C16

:082

914

2520

1890

1726

02

554

876

1044

1300

452

7018

93

(534

–93

8)(2

060

–35

70)

(148

0–

2310

)(1

50–

312)

(41

–69

)(6

06–

1150

)(7

80–

1190

)(3

710

–69

50)

(143

5–

2352

)

Cis

o16

:03

21

47

(2–

3)

Sum

C12

:0–

1200

3820

2460

424

142

1512

8014

1719

5074

4022

00

C16

:0(3

050

–53

30)

(205

0–

2880

)(2

58–

558)

(115

–16

8)(7

80–

1720

)(1

076

–16

00)

(524

0–

1000

0)(1

680

–27

30)

C17

:02

6741

75

21

428

4010

93

6185

(63

–72

)(4

–7)

(1–

1)(1

6–

40)

(24

–48

)(4

7–

80)



Tab

le8

.(C

onti

nued

) Co

wa

Bu

ffal

ob

n=

Bu

ffal

ob

Yak

c

(n=

1,ra

ng

eg

iven

wh

ere

n=

2)M

ared

n=

Mar

edD

on

keye

n=

Do

nke

yeD

rom

edar

yf

n=

Dro

med

aryf

cam

el

Bac

tria

ng

cam

el(n

=3

anim

als)

Llam

ahRe

ind

eeri

n=

Rein

dee

ri

Mo

ose

j

(n=

1,ra

ng

eg

iven

wh

ere

n=

2)

Cis

o17

:04

42

45

(3–

5)(2

8–

63)

C18

:036

514

952

964

1313

25

434

678

143

74

2120

1044

(220

–42

8)(5

48–

1237

)(7

51–

1180

)(8

–20

)(4

–7)

(299

–40

1)(7

24–

869)

(138

0–

2740

)

Cis

o18

:02

8(6

–10

)

C19

:01

19

C20

:03

832

14

1750

151

239

(73

–97

)(0

–1)

(14

–20

)

C21

:01

13

C22

:016

2Tr

25

413

118

(3–

7)

C23

:02

2(1

–3)

Tota

lSFA

∗18

7058

1035

4055

031

020

8022

3729

4011

240

4540

(426

0–

8810

)(2

920

–41

80)

(320

–75

0)(2

40–

380)

(151

0–

2600

)(1

849

–25

17)

(751

0–

1507

0)(3

110

–59

60)

Tota

lC4

–C

10∗

300

888

5910

115

937

540

514

8196

7

(535

–20

67)

(46

–15

9)(1

19–

200)

(369

–38

2)(7

29–

2098

)(7

41–

1193

)

Mo

no

un

satu

rate

dFA

C8

:11

145

C10

:13

1413

152

111

522

(12

–19

)(7

–22

)(8

–13

)

C12

:112

22

11

5541

(1–

4)(1

–2)

C13

:11

10

C14

:18

5519

137

25

340

1717

71

4354

(38

–81

)(4

–9)

(1–

8)(8

–61

)(1

2–

26)

C15

:14

3(3

–5)

C16

:111

147

613

723

292

171

296

298

(74

–25

2)(6

1–

91)

(171

–36

2)(1

07–

219)

(239

–35

6)

C16

:1n

-75

72

111

335

(5–

8)(9

–14

)

C16

:1ci

s-9

570

(55

–82

)

C17

:13

256

62

1

(22

–28

)(1

–11

)(1

–2)

C17

:1n

-81

14

C18

:181

214

1824

1101

1723

02

464

622

1119

733

424

9020

64

(501

–92

8)(1

131

–22

61)

(60

–38

5)(3

5–

58)

(381

–86

4)(7

61–

1424

)(1

720

–29

30)

(175

5–

2372

)



Tab

le8

.(C

onti

nued

) Co

wa

Bu

ffal

ob

n=

Bu

ffal

ob

Yak

c

(n=

1,ra

ng

eg

iven

wh

ere

n=

2)M

ared

n=

Mar

edD

on

keye

n=

Do

nke

yeD

rom

edar

yf

n=

Dro

med

aryf

cam

el

Bac

tria

ng

cam

el(n

=3

anim

als)

Llam

ahRe

ind

eeri

n=

Rein

dee

ri

Mo

ose

j

(n=

1,ra

ng

eg

iven

wh

ere

n=

2)

C18

:1ci

sn

-11

310

45

161

106

(78

–13

1)(1

2–

21)

C18

:1ci

sn

-12

137

C18

:1ci

sn

-13

114

C19

:11

46

C20

:19

4(1

–5)

1494

C20

:1n

-71

16

C20

:1n

-91

14

20:1

n-1

12

2(1

–2)

C21

:11

95

C22

:12

36(1

9–

53)

27

C24

:17

Tota

lMU

FA∗

812

2170

1180

432

7722

0013

0712

6027

3025

40

(136

0–

2770

)(2

10–

643)

(56

–99

)(1

787

–25

50)

(894

–16

57)

(196

0–

3170

)

Poly

un

satu

rate

dFA

C16

:210

9

C16

:345

C16

:425

C18

:212

08

9685

812

22

108

5997

7

(75

–13

9)(7

2–

106)

(58

–20

0)(9

7–

118)

(281

–16

73)

C18

:29-

c,12

-c3

93(9

0–

96)

C18

:375

952

613

184

(129

–25

6)2

8021

4

(11

–10

4)(4

5–

115)

(121

–30

6)

C18

:36-

9-12

-c1

1

C18

:39-

12-1

5-c

357

(26

–74

)

C20

:25

3(2

–4)

11

C20

:449

C20

:530

C22

:446

C22

:55

C22

:65



Tab

le8

.(C

onti

nued

)

Co

wa

Bu

ffal

ob

n=

Bu

ffal

ob

Yak

c

(n=

1,ra

ng

eg

iven

wh

ere

n=

2)M

ared

n=

Mar

edD

on

keye

n=

Do

nke

yeD

rom

edar

yf

n=

Dro

med

aryf

cam

el

Bac

tria

ng

cam

el(n

=3

anim

als)

Llam

ahRe

ind

eeri

n=

Rein

dee

ri

Mo

ose

j

(n=

1,ra

ng

eg

iven

wh

ere

n=

2)

n-3

C18

:32

5214

942

301

5659

153

n-3

(ALA

)(5

0–

54)

(56

–29

9)(2

3–

38)

(52

–66

)

C18

:4n

-32

1(1

–1)

C20

:3n

-35

3(1

–7)

1Tr

(0.4

)

20:4

n-3

1Tr

(0.3

)

C20

:5n

-3(E

PA)

26

(5–

7)Tr

(0.3

)2

Tr(T

r–2)

113

C22

:5n

-34

1(1

–1)

1Tr

(0.4

)2

12(9

–16

)

22:6

n-3

(DH

A)

Tr(0

.4)

21

(1–

2)

Tota

ln-3

∗58

9934

6959

65

(55

–61

)(5

8–

307)

(52

–66

(62

–69

)

n-6

C18

:2,n

-69

145

239

120

744

326

(LA

)(5

1–

241)

(29

–49

)(5

9–

82(1

22–

443)

C20

:2n

-64

5(4

–6)

22

(1–

2)

20:3

n-6

81

(1–

1)

C20

:4n

-62

6(6

–7)

92

(1–

6)2

31(2

5–

37)

C18

:3n

-62

1(T

r–1)

214

4(1

0–

279)

C22

:6n

-62

4(4

–5)

Tota

ln-6

∗10

153

4220

7450

1

(10

–12

)(5

7–

254)

(31

–52

)(5

9–

82(1

57–

759)

Tota

lPU

FA∗

195

454

(265

–45

4)14

656

0(3

04–

1021

)77

277

153

150

566

1460

(56

–96

)(2

31–

322)

(126

–16

9)(2

19–

828)

(671

–22

48)

tran

sFA

C16

:1tr

ans

216

(11

–22

)

C18

:1tr

ans

217

7

(141

–21

2)

C18

:1tr

ans

n-9

315

(14

–15

)1

53

C18

:1tr

ans

n-1

13

210

150

215

1

(159

–27

4)(1

30–

172)



Tab

le8

.(C

onti

nued

)

Co

wa

Bu

ffal

ob

n=

Bu

ffal

ob

Yak

c

(n=

1,ra

ng

eg

iven

wh

ere

n=

2)M

ared

n=

Mar

edD

on

keye

n=

Do

nke

yeD

rom

edar

yf

n=

Dro

med

aryf

cam

el

Bac

tria

ng

cam

el(n

=3

anim

als)

Llam

ahRe

ind

eeri

n=

Rein

dee

ri

Mo

ose

j

(n=

1,ra

ng

eg

iven

wh

ere

n=

2)

C18

:1tr

ans

n-4

18

C18

:1tr

ans

n-5

–6

133

C18

:1tr

ans

n-7

–8

159

C18

:1tr

ans

n-1

01

65

C18

:1tr

ans

n-1

21

59

C18

:1tr

ans

n-1

51

33

C18

:2t9

t12

tran

s2

28(2

2–

33)

C18

:2tr

ans

n-6

C18

:2c9

,t11

669

165

1(T

r–1)

2132

(n-6

),C

LA(4

0–

131)

(16

–26

)

C18

:210

-t,1

2-c

613

(5–

20)

tota

ltra

nsFA

∗24

528

161

(Tr–

1)21

182

461

(392

–70

7)(1

6–

26(4

40–

482)

tota

lCLA

(C18

:29-

c,11

-t)+

(C18

:210

-t,

12-c

)

1282

(45

–15

1)16

1(T

r–1)

32

Fro

mp

aper

s

Tota

lsat

ura

ted

951

6051

508

576

337

219

3029

50

(458

0–

6570

)(4

50–

693)

(335

–40

5)

Tota

lun

satu

rate

d3

1830

2750

469

313

28

(138

0–

2690

)(5

67–

773)

Tota

lMU

FAp

aper

s3

1960

537

83

130

142

(176

0–

2150

)(3

31–

465)

(95

–15

7)

Tota

lPU

FA3

204

529

83

151

182

pap

ers

(193

–21

6)(2

39–

405)

(100

–20

8)



Tab

le8

.(C

onti

nued

)

Co

wa

Bu

ffal

ob

n=

Bu

ffal

ob

Yak

c

(n=

1,ra

ng

eg

iven

wh

ere

n=

2)M

ared

n=

Mar

edD

on

keye

n=

Do

nke

yeD

rom

edar

yf

n=

Dro

med

aryf

cam

el

Bac

tria

ng

cam

el(n

=3

anim

als)

Llam

ahRe

ind

eeri

n=

Rein

dee

ri

Mo

ose

j

(n=

1,ra

ng

eg

iven

wh

ere

n=

2)

Tota

ltra

ns3

220

pap

ers

(174

–26

7)

tota

lCLA

(C18

:213

805

1

9-c,

11-t

)+(C

(44

–15

0)(T

r–1)

18:2

10-t

,12-

c)

n-3

tota

l6

452

94

(10

–10

9)(6

3–

124)

n-6

tota

l6

112

283

(78

–12

3)

∗ Cal

cula

ted

asth

esu

mo

frep

ort

edin

div

idu

alFA

.No

teth

atn

ot

allt

he

ind

ivid

ual

FAar

ere

po

rted

inth

ep

aper

s,as

som

ep

aper

so

nly

rep

ort

edva

lues

for

tota

lFA

.Th

eca

lcu

late

dva

lues

for

tota

lMU

FAan

dPU

FAd

on

ot

incl

ud

etr

ans

fatt

yac

ids,

wh

ich

are

giv

ense

par

atel

y.∗∗

Val

ues

fort

ota

lFA

asre

po

rted

inp

aper

s.Tr

,tra

ce=

less

than

1m

g.

SFA

,sat

ura

ted

fatt

yac

ids;

MU

FA,m

on

ou

nsa

tura

ted

fatt

yac

ids;

PUFA

,po

lyu

nsa

tura

ted

fatt

yac

ids;

TFA

,tra

nsfa

tty

acid

s;LA

,lin

ole

icac

id;A

LA,α

-lin

ole

nic

acid

;EPA

,eic

osa

pen

taen

oic

acid

;DH

A,d

oco

sah

exae

no

icac

id;

CLA

,co

nju

gat

edlin

ole

icac

id.

aC

ow

.45

bB

uff

alo

.Ave

rag

eva

lues

,usi

ng

valu

esfr

om

Nili

Ravi

and

Ku

nd

ibre

ed,41

Mu

rrah

bre

ed,d

ata

fro

mth

ree

diff

eren

tfe

eds)

,70–

72M

urr

ahb

reed

,dat

afr

om

earl

y,m

id,l

ate

lact

atio

n.13

4

cY

ak.A

vera

ge

valu

es,u

sin

gva

lues

fro

mK

hai

nag

bre

ed11

3an

dM

aiw

ab

reed

.37

dM

are.

Ave

rag

eva

lues

,usi

ng

valu

esfr

om

pri

mit

ive

Ko

nik

ho

rse,

85W

ielk

op

ols

kam

are

bre

ed,74

fou

rd

iffer

ent

lact

atio

nst

ages

,Hafl

ing

erb

reed

,fo

ur

diff

eren

tla

ctat

ion

stag

es,76

Ko

nik

Pols

ki,W

ielk

op

ols

kam

are

bre

edan

dPo

lish

Co

ld-b

loo

ded

Ho

rse

bre

eds,

49Ra

pid

Hea

vyD

raft

bre

edan

d1s

tan

d2n

dm

ilkco

llect

ion

dat

afo

rHafl

ing

eran

dQ

uar

terH

ors

eb

reed

s.77

eD

on

key.

Ave

rag

eva

lues

,usi

ng

valu

esfr

om

Rag

usa

na

bre

ed,8

Mar

tin

aFr

anca

bre

ed,t

wo

diff

eren

tfe

eds14

5an

dm

ilkfr

om

am

ixo

fRag

usa

na

and

Mar

tin

aFr

anca

bre

eds.

81

fD

rom

edar

yca

mel

.Ave

rag

eva

lues

,usi

ng

valu

esfr

om

Naj

dib

reed

78,1

46an

dM

ajah

eem

bre

ed.14

7

gLl

ama.

50

hRe

ind

eer53

(fo

urd

iffer

ent

lact

atio

nst

ages

).iM

oo

se52

(ave

rag

efo

ran

imal

s1

–3,

and

dat

afr

om

ref.

9ci

ted

ther

ein

).jB

actr

ian

cam

el60

(dat

afo

rth

ree

do

mes

tica

ted

cam

els)

.



Tab

le9

.Fa

tty

acid

con

ten

t,g

100

g−1

tota

lFA

(ave

rag

e,w

ith

ran

ge

inp

aren

thes

es)

Fatt

yac

idin

100

gto

talF

AC

ow

aB

uff

alo

bY

akc

Mar

edD

on

keye

Dro

med

aryf

Bac

tria

n†g

Llam

ah(n

=1)

Rein

dee

riM

oo

sej

aver

age†

Tota

lfat

(g10

0g

−1m

ilk)

3.3

7.4

6.8

1.6

0.7

3.0

5.0

4.2

16.1

8.6

(5.3

–9.

0)(5

.6–

9.5)

(0.5

–4.

2)(0

.3–

1.8)

(2.0

–6.

0)(4

.3–

5.7)

(2.7

–4.

7)(1

0.2

–21

.5)

(7.0

–10

.0)

Tota

lC4

–C

109.

613

0.9∗

6.7∗

23∗

13∗

9∗11

∗4∗

(8–

30)

(17

–25

)(5

–16

)(0

–15

)

Sum

C12

:0–

C16

:038

.555

47∗

28∗

21∗

43∗

30∗

45∗

56∗

27∗

(44

–76

)(3

9–

55)

(17

–37

)(1

7–

29)

(24

–54

)(2

3–

34)

(35

–75

)(2

1–

33)

Tota

lsat

ura

ted

60.0

7465

3857

6047

∗65

84∗

53

(65

–94

)(3

0–

46)

(49

–60

)(3

9–

53)

(56

–11

3)(5

2–

58)∗

Tota

lun

satu

rate

d26

3546

4235

(20

–38

)(3

8–

51)

Tota

lMU

FA26

.028

18∗

2520

73∗

28∗

3120

∗31

∗

(25

–31

)(2

2–

31)

(14

–23

)(5

6–

80)

(19

–35

)(1

5–

24)

Tota

ln-3

17∗

122.

4∗[0

.5∗ ]

(0–

2)(4

–20

)(9

–18

)(0

–1)

Tota

ln-6

210

∗11

1∗4∗

8

(1–

2)(4

–17

)(1

–6)

(4–

21)

Tota

lPU

FA6.

33

2∗20

2310

∗3∗

44∗

14∗

(16

–27

)(1

5–

31)

(7–

10)

(3–

4)(2

–6)

(8–

25)

Tota

ltra

nsFA

30.

2∗[0

.1∗ ]

33∗

(3–

4)

Tota

lCLA

(C18

:29-

c,11

-t)+

(C18

:210

-t,1

2-c)

1[0

.2∗ ]

[0.1

][0

.4∗ ]

1

(1–

2)(0

–1)

C18

:3n

-3(A

LA)

16.

24

21

[0.4

]

(4–

20)

(3–

6)

C20

:5n

-3(E

PA)

[0.0

2][0

.13]

[0.5

]

C22

:6n

-3(D

HA

)[0

.03]

[0.1

5]

C18

:2,n

-6(L

A)

106

12

28

(3–

16)

(4–

7)(1

–2)

(1–

3)(4

–21

)

∗ Den

ote

sva

lues

that

wer

eca

lcu

late

das

the

sum

oft

he

rep

ort

edin

div

idu

alva

lues

.Val

ues

less

than

0.1%

are

insq

uar

eb

rack

ets.

Cal

cula

ted

valu

esfo

rPU

FAan

dM

UFA

do

no

tin

clu

de

tran

sFA

,wh

ich

are

inth

eir

ow

nca

teg

ory

.Wh

enth

e%

FAh

adto

be

calc

ula

ted

,th

eav

erag

eva

lue

for

tota

lfat

ob

tain

edfr

om

the

pap

ers

rep

ort

ing

the

FAw

asg

ener

ally

use

d(r

ath

erth

anth

eav

erag

efo

rth

esp

ecie

s)in

the

calc

ula

tio

ns.

Tota

lFA

do

esn

ot

equ

al10

0%b

ecau

sen

ot

allp

aper

sre

po

rted

valu

esfo

rall

FA.

†Fo

rB

actr

ian

cam

elan

dm

oo

sem

ilk,v

ery

littl

ed

ata

wer

eav

aila

ble

,an

dla

rge

diff

eren

ces

wer

eo

bse

rved

bet

wee

nan

imal

sw

ith

inth

esa

me

stu

dy.

Ther

efo

re,t

he

dat

aw

ere

con

sid

ered

atan

imal

leve

l:th

eav

erag

efo

rB

actr

ian

cam

eld

ata

are

fro

mth

ree

(do

mes

tica

ted

)an

imal

s,re

po

rted

inJi

rim

utu

;60m

oo

sed

ata

are

fort

hre

ean

imal

sre

po

rted

inC

oo

kci

ted

ther

ein

.52

aC

ow

.45

bB

uff

alo

.Ave

rag

eva

lues

,usi

ng

valu

esfr

om

Nili

Ravi

bre

ed,41

Mu

rrah

bre

ed,d

ata

fro

mth

ree

diff

eren

tfe

eds,

70–

72M

urr

ahb

reed

,dat

afr

om

earl

y,m

id,l

ate

lact

atio

n.13

4

cY

ak.A

vera

ge

valu

es,u

sin

gva

lues

fro

mK

hai

nag

bre

ed11

3an

dM

aiw

ab

reed

.37

dM

are.

Ave

rag

eva

lues

,usi

ng

valu

esfr

om

pri

mit

ive

Ko

nik

ho

rse,

85W

ielk

op

ols

kam

are

bre

ed,f

ou

rd

iffer

ent

lact

atio

nst

ages

,74H

aflin

ger

bre

ed,f

ou

rd

iffer

ent

lact

atio

nst

ages

,76K

on

ikPo

lski

,Wie

lko

po

lska

mar

eb

reed

and

Polis

hC

old

-blo

od

edH

ors

eb

reed

s,49

Rap

idH

eavy

Dra

ftb

reed

and

1st

and

2nd

milk

colle

ctio

nd

ata

forH

aflin

ger

and

Qu

arte

rHo

rse

bre

eds.

77

eD

on

key.

Ave

rag

eva

lues

,usi

ng

valu

esfr

om

Rag

usa

na

bre

ed,8

Mar

tin

aFr

anca

bre

ed,t

wo

diff

eren

tfe

eds14

5an

dm

ixo

fRag

usa

na

and

Mar

tin

aFr

anca

bre

eds.

81

fD

rom

edar

yca

mel

.Ave

rag

eva

lues

,usi

ng

valu

esfr

om

Naj

dib

reed

78,1

46an

dM

ajah

eem

bre

ed.14

7

gLl

ama.

50

hRe

ind

eer.

Dat

afr

om

fou

rdiff

eren

tla

ctat

ion

stag

es.53

iM

oo

se.52

Ave

rag

efo

ran

imal

s1

–3,

and

dat

afr

om

ref.

9ci

ted

ther

ein

.jB

actr

ian

cam

el.D

ata

fort

hre

ed

om

esti

cate

dca

mel

s.60



Tab

le1

0.

Bu

ffal

om

ilkco

mp

osi

tio

na

Per1

00g

milk

Ave

rag

e±

SDRa

ng

eB

reed

and

loca

tio

nw

ith

low

est

valu

eRe

f.fo

rlo

wes

tva

lueb

Bre

edan

dlo

cati

on

wit

hh

igh

est

valu

eRe

f.fo

rhig

hes

tva

lueb

Wat

er(g

)83

.2±

0.6

82.3

–84

.0M

urr

ah×

Med

iter

ran

ean

(cro

ss-b

reed

)A

vera

ge96

,107

Bu

lgar

ian

Bu

ffal

o×

Mu

rrah

bre

ed(B

ulg

aria

)

97

n=

11,n

′ =51

Tota

lso

lids

(g)

17.0

±0.

616

.1–

17.7

Tod

a(In

dia

)98

Mu

rrah

×M

edit

erra

nea

n(c

ross

-bre

ed)

(Arg

enti

na,

Ger

man

y)

96,1

07

n=

8,n′ =

32

Solid

s–

no

tfa

t(g

)9.

7±

0.8

8.3

–11

.3N

on

-des

crip

th

illb

uff

alo

10K

utt

anad

Dw

arfB

uff

alo

(Ind

ia,K

eral

a)10

6

n=

9,n′ =

40(In

dia

/Ku

mao

nre

gio

n)

Pro

tein

,to

tal(

g)

4.0

±0.

52.

7–

4.6

No

n-d

escr

ipt

hill

bu

ffal

o10

Med

iter

ran

ean

bu

ffal

oA

vera

ge10

8,10

9

n=

10,n

′ =42

(Ind

ia/K

um

aon

reg

ion

)

Fat,

tota

l(g

)7.

4±

0.9

5.3

–9.

0B

ulg

aria

nB

uff

alo

×M

urr

ahb

reed

97B

had

awar

iA

vera

ge98

n=

18,n

′ =75

(Bu

lgar

ia)

Lact

ose

(g)

4.4

±0.

63.

2–

4.9

Ku

ttan

adD

war

fBu

ffal

o(In

dia

,Ker

ala)

106

Bu

lgar

ian

Mu

rrah

Ave

rag

e97,1

04,1

05

n=

6,n′ =

23

Ash

(g)

0.8

±0.

040.

7–

0.8

Meh

san

a95

Mu

rrah

×M

edit

erra

nea

n(c

ross

-bre

ed)

96

n=

5,n′ =

12(In

dia

,Bo

mb

ay)

(Arg

enti

na)

Cal

ciu

m(m

g)

191

±38

147

–22

0K

utt

anad

Dw

arfB

uff

alo

(Ind

ia/K

eral

a)10

6Eg

ypti

anb

uff

alo

57

n=

5,n′ =

9

Mag

nes

ium

(mg

)12

±5

2–

16K

utt

anad

Dw

arfB

uff

alo

(Ind

ia/K

eral

a)10

6M

urr

ah(In

dia

–B

om

bay

;Fra

nce

)95

,129

n=

5,n′ =

6

Pho

sph

oru

s(m

g)

185

±94

102

–29

3Eg

ypti

anb

uff

alo

57M

ehsa

na

(Ind

ia,B

om

bay

)95

n=

5,n′ =

8

Cit

ric

acid

(mg

)21

4±

1918

5–

224

Mu

rrah

(Ind

ia–

Bo

mb

ay;F

ran

ce)

95,1

29Eg

ypti

anb

uff

alo

57

n=

4,n′ =

5

Ch

ole

ster

ol(

mg

)8

±3

4–

10Eg

ypti

anb

uff

alo

57N

iliRa

vi(P

akis

tan

)41

n=

3,n′ =

3

Pho

sph

olip

ids,

tota

l(m

g)

20Eg

ypti

anb

uff

alo

57

n=

1,n′ =

1

aO

nly

pro

xim

ates

and

dat

aw

ith

n≥

4ar

esh

ow

n,w

ith

the

exce

pti

on

ofa

few

nu

trie

nts

wh

ich

hav

eb

een

incl

ud

edb

ecau

seth

eyd

on

ot

bel

on

gin

any

oft

he

tab

les

forn

utr

ien

tg

rou

ps.

bW

her

ed

ata

for

the

sam

eb

reed

wer

eav

aila

ble

fro

mm

ore

than

on

est

ud

y,th

em

ean

valu

efo

rth

eb

reed

was

calc

ula

ted

and

use

d,a

nd

the

refe

ren

ces

fro

mw

her

eth

ein

div

idu

alva

lues

wer

eta

ken

are

giv

en.T

he

nu

mb

ero

fen

trie

sin

the

smal

lerd

atab

ase

thu

so

bta

ined

isin

dic

ated

by

n.Th

en

um

ber

ofd

ata

po

ints

bef

ore

aver

agin

gfo

rbre

edis

ind

icat

edb

yn′ .



Tab

le1

1.

Yak

milk

com

po

siti

on

a

Per1

00g

milk

Ave

rag

e±

SDRa

ng

eB

reed

and

loca

tio

nw

ith

low

est

valu

eRe

f.fo

rlo

wes

tva

lueb

Bre

edan

dLo

cati

on

wit

hh

igh

est

valu

eRe

ffo

rhig

hes

tva

lueb

Ener

gy

(kJ)

368

349

–38

2Ti

anzh

uW

hit

eA

vera

ge

forv

alu

esfo

rear

ly,m

idan

dla

tela

ctat

ion

117

(Ch

ina)

11

(n=

2,n′ =

4)(C

hin

a/G

ansu

pro

vin

ce)

Ener

gy

(kca

l)89

87–

91Ti

anzh

uW

hit

eA

vera

ge

forv

alu

esfo

rear

ly,m

idan

dla

tela

ctat

ion

117

(Ch

ina)

11

(n=

2,n′ =

4)(C

hin

a/G

ansu

pro

vin

ce)

Wat

er(g

)82

.6±

1.8

75.3

–84

.4Y

ak(M

on

go

lia)

143

(Ind

ia)

110

(n=

23,n

′ =27

)

Tota

lso

lids

(g)

17.2

±0.

915

.6–

18.8

Kh

ain

ag(y

akh

ybri

d)(

Mo

ng

olia

)11

3(In

dia

)11

1

(n=

21,n

′ =24

)

Solid

s–

no

tfa

t(g

)9.

9±

1.7

7.1

–11

.5(In

dia

)11

0(In

dia

)11

1

(n=

7,n′ =

9)

Pro

tein

,to

tal(

g)

5.2

±0.

54.

2–

5.9

Mai

wa

37,1

20(In

dia

/Pra

des

h)

118

(n=

26,n

′ =32

)

Fat,

tota

l(g

)6.

8±

0.9

5.6

–9.

5K

hai

nag

113

Pam

ir11

9

(n=

41,n

′ =54

)(M

on

go

lia)

Lact

ose

(g)

4.8

±0.

63.

3–

6.2

Sib

u(T

ibet

)12

0(M

on

go

lian

)14

3

(n=

28,n

′ =34

)

Ash

(g)

0.8

±0.

10.

4–

1.0

Yak

115

Jial

i11

7,12

0

(n=

23,n

′ =28

)

Cal

ciu

m(m

g)

129

±7

119

–13

4(In

dia

)11

1K

hai

nag

113

(n=

54,n

′ =5)

(Mo

ng

olia

)

Lact

icac

id(m

g)

170

(Tib

et)

114

(n′ =

1,n

=1)

β-L

acto

glo

bu

lin(m

g)

708

±24

653

4–

882

Mai

wa

37Ji

ulo

ng

Ave

rag

eo

fval

ues

forfi

rsts

um

mer

seas

on

afte

rca

lvin

gan

dse

con

dsu

mm

erse

aso

naf

ter

calv

ing

wit

ho

ut

calv

ing

agai

n(h

alfm

ilk)11

5

(n=

3,n′ =

4)(C

hin

a)

α-L

acta

lbu

min

(mg

)19

8±

819

3–

204

Mai

wa

37Ji

ulo

ng

Ave

rag

eo

fval

ues

for:

first

sum

mer

seas

on

afte

rcal

vin

gan

dse

con

dsu

mm

erse

aso

naf

terc

alvi

ng

wit

ho

ut

calv

ing

agai

n(h

alf

milk

)115

(n=

3,n′ =

4)(C

hin

a)

Seru

mal

bu

min

(mg

)10

7±

3979

–13

4Ji

ulo

ng

Ave

rag

eo

fval

ues

for:

first

sum

mer

seas

on

afte

rcal

vin

gan

dse

con

dsu

mm

erse

aso

naf

terc

alvi

ng

wit

ho

ut

calv

ing

agai

n(h

alf

milk

)115

Mai

wa

(Ch

ina)

37

(n=

3,n′ =

4)

aO

nly

pro

xim

ates

and

dat

aw

ith

n≥

4ar

esh

ow

n,w

ith

the

exce

pti

on

ofa

few

nu

trie

nts

wh

ich

hav

eb

een

incl

ud

edb

ecau

seth

eyd

on

ot

bel

on

gin

any

oft

he

tab

les

forn

utr

ien

tg

rou

ps.

bW

her

ed

ata

fort

he

sam

eb

reed

wer

eav

aila

ble

fro

mm

ore

than

on

est

ud

y,th

em

ean

valu

efo

rth

eb

reed

was

calc

ula

ted

and

use

d,a

nd

the

refe

ren

ces

fro

mw

her

eth

ein

div

idu

alva

lues

wer

eta

ken

are

giv

en.T

he

nu

mb

ero

fen

trie

sin

the

smal

lerd

atab

ase

thu

so

bta

ined

isin

dic

ated

by

n.Th

en

um

ber

ofd

ata

po

ints

bef

ore

aver

agin

gfo

rbre

edis

ind

icat

edb

yn′ .



Tab

le1

2.

Mar

em

ilkco

mp

osi

tio

na

Per1

00g

milk

Ave

rag

e±

SDRa

ng

eB

reed

wit

hlo

wes

tva

lue

Ref.

forl

ow

est

valu

ebB

reed

wit

hh

igh

est

Ref.

forh

igh

est

valu

eb

Ener

gy

(kJ)

193

±24

177

–21

0H

aflin

ger

Lact

atio

nav

erag

e92Pr

zew

alsk

iho

rse

83

n=

2,n′ =

6

Ener

gy

(kca

l)46

±6

42–

50H

aflin

ger

Lact

atio

nav

erag

e92Pr

zew

alsk

iho

rse

83

n=

2,n′ =

6

Wat

er(g

)89

.8±

0.8

87.9

–91

.3Sa

dd

lep

on

y12

3Lu

sita

no

Lact

atio

nav

erag

e94

n=

13,n

′ =30

Tota

lso

lids

(g)

10.5

±0.

69.

4–

12.1

Hafl

ing

erA

vera

ge

ofm

ilkfr

om

1st

and

2nd

colle

ctio

ns77

,82

Sad

dle

po

ny

123

n=

19,n

′ =30

Pro

tein

,to

tal(

g)

2.0

±0.

41.

4–

3.2

San

am

ares

(mts

yri)

Ave

rag

eva

lue57

Palo

min

o13

1

n=

20,n

′ =33

Fat,

tota

l(g

)1.

6±

(0.7

)0.

5–

Lusi

tan

oLa

ctat

ion

aver

age93

,94

Sad

dle

po

ny

123

n=

23,n

′ =45

4.2

Lact

ose

(g)

6.6

±(0

.4)

5.6

–7.

2B

ury

atA

vera

ge

valu

e57Tr

ott

ers

Ave

rag

eva

lue57

n=

16,n

′ =31

Ash

(g)

0.4

±(0

.1)

0.3

–0.

5Im

pro

ved

Kir

gh

izA

vera

ge

valu

e57B

ury

atA

vera

ge

valu

e57

n=

18,n

′ =38

Cal

ciu

m(m

g)

95±

(19)

76–

124

Tho

rou

gh

bre

dLa

ctat

ion

aver

age12

4,13

8Pa

lom

ino

131

n=

9,n′ =

26

Iro

n(m

g)

0.10

±(0

.05)

0.03

–0.

15Th

oro

ug

hb

red

Lact

atio

nav

erag

e124,

138

Ital

ian

sad

dle

ho

rse

122,

135

n=

4,n′ =

6

Mag

nes

ium

(mg

)7

±(2

)4

–12

Lusi

tan

oLa

ctat

ion

aver

age94

Palo

min

o13

1

n=

8,n′ =

1858

±(1

3)

Pho

sph

oru

s(m

g)

n=

8,n′ =

2043

–83

Prze

wal

skih

ors

eLa

tela

ctat

ion

83,1

24Pa

lom

ino

131

Pota

ssiu

m(m

g)

51±

(20)

25–

87Sh

etla

nd

po

ny

124,

138

Palo

min

o13

1

n=

9,n′ =

18

Sod

ium

(mg

)16

±(3

)13

–20

Shet

lan

dp

on

y12

4,13

8H

aflin

ger

137

n=

6,n′ =

10

Zin

c(m

g)

0.2

±(0

.1)

0.2

–0.

3Sh

etla

nd

124,

138

Ital

ian

sad

dle

ho

rse

122,

135



Tab

le1

2.

(Con

tinu

ed)

Per1

00g

milk

Ave

rag

e±

SDRa

ng

eB

reed

wit

hlo

wes

tva

lue

Ref.

forl

ow

est

valu

ebB

reed

wit

hh

igh

est

Ref.

forh

igh

est

valu

eb

n=

5,n′ =

8Po

ny

Co

pp

er(m

g)

0.05

±(0

.04)

0.02

–0.

11Pr

zew

alsk

iho

rse

Late

lact

atio

n83

,124

Bar

dig

ian

o12

2

n=

5,n′ =

8

Vit

amin

C(m

g)

4.3

±(3

.3)

1.7

–8.

1Sa

dd

lep

on

y12

3Pa

lom

ino

131

n=

3,n′ =

6

Thia

min

e(m

g)

0.03

±(0

.01)

0.02

–0.

04Sa

dd

lep

on

y12

3Pe

rch

ero

nA

vera

ge

fro

mth

ree

entr

ies12

3

n=

2,n′ =

4

Rib

ofla

vin

(mg

)0.

020.

011

–0.

026

Perc

her

on

Ave

rag

efr

om

thre

een

trie

s123

Sad

dle

po

ny

123

n=

2,n′ =

4

Nia

cin

(mg

)0.

07Sa

dd

lep

on

y;A

vera

ge

fro

m(t

hre

een

trie

s123

)8

n=

2,n′ =

4Pe

rch

ero

n

Cas

ein

(mg

)85

5±

3083

5–

890

Hafl

ing

erA

vera

ge

ofm

ilkfr

om

1st

and

2nd

colle

ctio

ns77

Qu

arte

rA

vera

ge

ofm

ilkfr

om

1st

and

2nd

colle

ctio

ns77

n=

3,n′ =

5

Wh

eyp

rote

in(m

g)

790

±14

966

5–

955

Hafl

ing

erA

vera

ge

ofm

ilkfr

om

1st

and

2nd

colle

ctio

ns77

Qu

arte

rA

vera

ge

ofm

ilkfr

om

1st

and

2nd

colle

ctio

ns77

n=

3,n′ =

5

Sial

icac

id4.

6m

gB

ard

igia

no

;Ita

lian

sad

dle

ho

rse

122

n=

2,n′ =

2

aO

nly

pro

xim

ates

and

dat

aw

ith

n≥

4ar

esh

ow

n,w

ith

the

exce

pti

on

ofa

few

nu

trie

nts

wh

ich

hav

eb

een

incl

ud

edb

ecau

seth

eyd

on

ot

bel

on

gin

any

oft

he

tab

les

forn

utr

ien

tg

rou

ps.

bW

her

ed

ata

for

the

sam

eb

reed

wer

eav

aila

ble

fro

mm

ore

than

on

est

ud

y,th

em

ean

valu

efo

rth

eb

reed

was

calc

ula

ted

and

use

d,a

nd

the

refe

ren

ces

fro

mw

her

eth

ein

div

idu

alva

lues

wer

eta

ken

are

giv

en.T

he

nu

mb

ero

fen

trie

sin

the

smal

lerd

atab

ase

thu

so

bta

ined

isin

dic

ated

by

n.Th

en

um

ber

ofd

ata

po

ints

bef

ore

aver

agin

gfo

rbre

edis

ind

icat

edb

yn′ .



content was high, with an average value of 73 g 100 g−1 totalFA. The milk also contained a small amount (2 g 100 g−1 totalFA) of ALA. The content of very short-chain fatty acids (C4–C8)was lowest in dromedary milk among all the species with valuesreported here. It has been suggested that these FA, which areproduced by cellulose fermentation in the rumen, may be rapidlymetabolized by camel tissue and are therefore not excreted in themilk.78

Only one study on Bactrian camel milk was available, andsince the variation in the fatty acid composition among the threeanimals was quite large, the ranges observed by the authors arealso reported here. A small amount (2 g 100 g−1 total FA) of LAwas reported in the milk.

One study50 reported the FA composition of llama milk. As g100 g−1 total FA, the proportions of SFA, C4–C10, MUFA and PUFAin llama milk were comparable to the values for cow milk. Themilk also contained trans FA (3 g/100 g−1 total FA, mainly C18 : 1trans-11), and a very small amount of CLA (0.4 g 100 g−1 total FA).

Cervid species (reindeer and moose). The FA profile of reindeermilk showed the highest amount of SFA (84 g 100 g−1 FA). Whenthe fatty acid content in 100 g milk was considered, the total SFAcontent of reindeer milk far exceeded that of other milks, havingan average of 11240 mg 100 g−1 milk (compared with 1870 mg100 g−1 cow milk). Calculated from the average value, two cupsof reindeer milk would contain 56. 2 g saturated fat. For a womanwho requires 2200 kcal per day this would provide 23% of the totalenergy intake. The population nutrient intake goals for preventingdiet-related chronic diseases recommend that saturated fat shouldaccount for less that 10% of the total energy intake.79

Moose milk FA data were limited, consisting of data for threeanimals reported in Cook et al.52 cited therein. According to thesestudies, moose milk had a high content of PUFA compared withcow milk.

Interbreed differences in milk compositionBreed differences are one of the factors known to affect milkcomposition.25 Before evaluating breed differences, it is importantto understand the normal biological variation within a breed. Apreliminary analysis of data for the Murrah breed of buffalo, whichprovided the greatest number of data points at the level of breed(up to 19 data points for some nutrients), was therefore carriedout (data not shown). These data represented different lactationperiods, feeds and seasons. Fairly small differences were presentin proximate composition (protein: 0.3 g 100 g−1; total fat: 0.8 g100 g−1; lactose: 0.4 g 100 g−1; solids-not-fat: 0.5 g 100 g−1). Incontrast, the differences among breeds for buffalo (Table 10) areconsiderably larger (protein: 1.9 g 100 g−1; total fat: 3.7 g 100 g−1;lactose: 1.7 g 100 g−1; solids-not-fat: 3 g 100 g−1). Therefore, adifference in nutrient content among different breeds which isnotably larger than the difference seen within a single breed mayat least be partially attributed to breed.

However, the standard deviations for individual fatty acids werevery large even within the Murrah breed. When the fatty acids werecalculated as per 100 g total FA, these large differences were stillpresent, indicating that many factors in addition to breed affectthe FA composition. This agrees with published data that showthat the FA composition is particularly prone to vary with factorsthat include feed, lactation period and season.24,26,27

Buffalo milkLarge differences were reported for fat values; Bulgarian buffalocross Murrah breed milk had the lowest reported content of5.3 g 100 g−1, while the Bhadawari buffalo milk had an averagefat content of 9 g 100 g−1. Protein values also varied, from 2.7 g100 g−1 in ‘hill’ buffalo milk to 4 g 100 g−1 in Mediterranean buffalomilk. Variations were also seen in mineral values (e.g. calcium,147–220 mg 100 g−1; phosphorus, 102–293 mg 100 g−1), but itis not clear if these differences can be attributed to differencesamong breeds. Individual references may be found in Table 10.

Yak and mithun milksThe total fat in yak milk ranged from 5.6 to 9.5 g 100 g−1 andaveraged 6.8 g 100 g−1, with milk from the Pamir breed having thehighest reported content and milk from the Khainag breed fromMongolia having the lowest. The lactose value of yak milk from theSibu breed from Tibet was nearly half that reported for a Mongolianyak (3.3 versus 6.2 g 100 g−1). Fairly large differences were alsoseen in the mineral values, but the small number of studies (oneto five studies per element) precludes any conclusions. Individualreferences may be found in Table 11.

Mare and donkey milksThe range for total fat in mare milk was 0.5–4.2 g 100 g−1, withan average value of 1.6 g 100 g−1 milk. The breed with the lowestvalue was Lusitano (average from two studies and a number oflactation periods). The value for the highest fat content (Saddlepony) was 2.6 times greater than the average fat content. Largedifferences between the lowest and highest concentrations wereapparent for some of the essential elements. For example, milkfrom the Palomino breed had 48 mg more calcium per 100 g milkthan milk from the Thoroughbred breed. The potassium contentdiffered by 62 mg 100 g−1 (25–87 mg 100 g−1) between the breedwith the lowest value, Shetland pony, and highest value, Palomino.Phosphorus content was again highest for milk from the Palominobreed (83 mg 100 g−1), being almost double the value for milkfrom the Przewalski horse (42 mg 100 g−1). Individual referencesmay be found in Table 12.

In donkey milk, the casein content varied from 612 mg 100 g−1

milk from the Jiangyue breed80 to 870 mg 100 g−1 milk for a mixedmilk from Ragusana and Martina Franca breeds.81

Dromedary camel, Bactrian camel, llama and alpaca milksThe total fat in dromedary camel milk ranged from 2 to 6 g100 g−1 milk, with the lowest fat content reported for milk fromthe Kachchhii breed39 and the highest for milk from the Arvanabreed.32 Fairly large interbreed differences were also reportedfor certain minerals. For example, manganese ranged from 60 to180 µg 100 g−1 milk: the Najdi breed milk showed a value threetimes higher than the value for milk from the Majaheim breed. Thedifference between the highest (Najdi breed) and lowest (Hamrabreed) potassium contents in dromedary milk was 49 mg (range124–173 mg 100 g−1 milk). Individual references may be found inTable 13.

Variation in milk composition with lactation stageReports suggest that stage of lactation is the most important factorto influence milk composition.82 Since a substantial number ofstudies looked at the effect of lactation stage on milk composition,the data from these studies are summarized below.



Table 13. Dromedary camel milk compositiona

Per 100 g milk Average ± SD RangeBreed with

lowest valueRef. for

lowest valuebBreed with

highest valueRef. for

highest valueb

Water (g) 89.0 ± 0.5 88.3–89.4 Majaheim 36 Hamra 125

n = 3, n′ = 3

Protein, total (g) 3.1 ± 0.5 2.4–4.2 Kachchhi 126 Wadah 125

n = 11, n′ = 12

Fat, total (g) 3.2 ± 1.1 2.0–6.0 Kachchhi 126 Arvana 32

n = 15, n′ = 23

Ash (g) 0.8 Wadah, Hamra,Majaheim, Bikaneri,Najdi, Turkana,Jaisalmeri, Kachchhiand Somali

36,38,39,63,87,125,126

n = 10, n′ = 12

Lactose (g) 4.3 ± 0.4 3.5–4.9 Arvana 32 Somali 87

n = 12, n′ = 15

Vitamin C (mg) 6.7 ± 7 2.5–18.4 Majaheem 63 Arvana 32

n = 5, n′ = 5

Calcium (mg) 114 ± 6 105–120 Arvana 32 Majaheem 125

n = 5, n′ = 5

Iron (mg) 0.21 0.17–0.26 Arvana 32 Hamra 125

n = 5, n′ = 5

Magnesium (mg) 13 ± 1 12–14 Hamra 125 Najdi 38

n = 4, n′ = 4

Phosphorus (mg) 86 ± 3 83–90 Arvana 32 Hamra 125

n = 5, n′ = 5

Potassium (mg) 151 ± 25 124–173 Hamra 125 Najdi 38

n = 4, n′ = 4

Sodium (mg) 66 ± 6 59–73 Najdi 38 Wadah 125

n = 4, n′ = 4

Zinc (mg) 0.6 ± 0.1 0.4–0.6 Najdi 38 Majaheim 125

n = 4, n′ = 4

Copper (mg) 0.15 0.12–0.17 Majaheim 125 Wadah 125

n = 4, n′ = 4

Manganese (µg) 106 60–180 Majaheim 125 Najdi 38

n = 4, n′ = 4

Free cholesterol (mg) 9 Najdi 78,146

n = 1, n′ = 4

Cholesteryl ester (mg) 5 Najdi 78,146

n = 1, n′ = 4

Total cholesterol (mg) 19 Najdi 78,146

n = 1, n′ = 3

a Only proximates and data with n ≥ 4 are shown, with the exception of a few nutrients which have been included because they do not belong inany of the tables for nutrient groups.b Where data for the same breed were available from more than one study, the mean value for the breed was calculated and used, and the referencesfrom where the individual values were taken are given. The number of entries in the smaller database thus obtained is indicated by n. The number ofdata points before averaging for breed is indicated by n′.

Protein

The milk from most species showed a pattern of being higher inearly and late lactation and lower in mid lactation. However, someequids have been reported to produce relatively dilute milk inmid to late lactation, e.g. Przewalski horse83 and Littoral-Dinaricdonkey,84 where a decrease in both milk yield and in proteinpercentage have been reported as lactation progresses. Wild andsemi-domestic ruminants are reported to give milk richer in bothprotein and fat, particularly in late lactation, than do domesticatedspecies, partially to compensate for the declining rate of milkintake by the calf during late lactation.53,55

Fat

The changes in fat were reported to be difficult to interpret, beingstrongly related to seasonal feeding effects.74,85,86 Especially incamel milk, the availability of feed and water in arid areas has beenshown to influence milk composition.87 The fat percentage wasshown to increase as lactation progressed for milk from buffalo,e.g. Murrah breed88,89 and Nili Ravi breed;90 mithun,46 llama20 andalpaca raised at low altitudes;91 and reindeer.55 In contrast, the fatpercentage decreased as lactation progressed in mare milk, e.g.Wielkopolska breed,74 Haflinger breed,92 Lusitano breed93,94 andprimitive Konik horse85 and donkey milk (Littoral-Dinaric breed).84



LactoseLactose was reported to be the least variable component in somemilk, e.g. Murrah buffalo,88 mithun48 donkey (Littoral-Dinaric)84

and Jiangyue breeds80 and in Bactrian camel,65 being osmoticallyactive and functioning as a regulator of water content in milk.55

The lactose percentage showed a slight increase as lactationprogressed in mare milk, e.g. Haflinger breed92 and Lusitanobreed,93,94 whereas in reindeer milk the lactose percentagedecreased significantly as lactation progressed.55

CONCLUSIONThis review attempts to widen the biodiversity knowledge baseby bringing together available data on the composition of milkfrom underutilized species and minor dairy animals at breed level.Using cow milk as a reference, comparisons of milk compositionwere made among species and between breeds within the samespecies. Significant differences were found both for macro- andmicronutrients, although the limitation or lack of data for somespecies impeded the analysis, making comparisons difficult. Manyfactors influence nutrient content in milk. Large interspeciesdifferences in nutrient composition were evident, ranging from 0.7to 16.1 g 100 g−1 for total fat, from 1.6 to 10.5 g 100 g−1 for proteinand from 2.6 to 6.6 g 100 g−1 for lactose, when species averageswere considered. Interbreed differences in fat and protein contentswere found in buffalo, mare, yak and dromedary camel, milks witha difference of ∼4 g fat and 2 g protein 100 g−1 milk betweenthe highest and lowest values. For mare, buffalo and dromedarycamel milks, large interbreed differences in mineral content wereapparent. The vitamin C content in dromedary camel milk rangedfrom 2.5 to 18.4 mg 100 g−1, which can mean the differencebetween nutritional adequacy and deficiency. Equine milk mayhave useful applications in infant feeding, due to its low proteinand mineral contents (lower renal solute load) and similarities inprotein and fat composition to those in human milk. Similar tohuman milk, equine milk has a high lactose content. However,further studies are needed, particularly because adverse effectson iron nutrition can be expected. The nutritional value of milkwould depend on the specific nutritional needs of the populationgroup or individual in question; better knowledge of interspeciesand interbreed differences would allow more informed choices fordiet interventions.

In addition to bringing together most of the available data forminor dairy animals and buffalo breeds, this study has helpedto identify areas where data are limited or inadequate either atbreed level or at species level, for example, vitamin data anddata on cholesterol for milk from most of the species covered inthe review. As milk composition is known to change because ofvarious factors in addition to breed differences, the importanceof reporting data on feeding regimes, lactation stages etc. whenpresenting dairy studies was also apparent. One limitation of thisstudy is that we concentrated largely on breed-level data: data thatwere reported only at species level were not collected (except fordata of underutilized species), which makes the species-level datacomparison less complete than it could have been. For example,studies on buffalo that reported values for generic buffalo milk(without specifying the breed) were excluded.

This review has shown there are large differences in mostnutrients in milk from different species and even among breedswithin the same species. Generating nutrient data in milk from lesscommonly used species of dairy animals and reporting nutritionaldata at the breed level will broaden the knowledge base on dairy

animal biodiversity as essential sources of nutrients, facilitate themaintenance of local species and breeds, and contribute towardsthe conservation of diverse genetic resources for improving humannutrition.

ACKNOWLEDGEMENTSThe authors are grateful to Jerome Mounsey for advice onanalysing the data and ideas for improving the manuscript. Manythanks to Maylis Razes for help with SPSS and Isabella McDonnellfor carefully reading the final draft.

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Composition of milk from minor dairy animals and buffalo breeds: a biodiversity perspective

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