DOI: 10.1111/j.1439-0396.2008.00832.
ORIGINAL ARTICLE
Skin surface lipids and skin and hair coat condition in dogs fed
increased total fat diets containing polyunsaturated fatty acids
N. A. Kirby, S. L. Hester, C. A. Rees, R. A. Kennis, D. L. Zoran and J. E. Bauer
Department of Veterinary Small Animal Medicine and Surgery and Faculty of Nutrition, College of Veterinary Medicine and Biomedical Sciences,
Texas A&M University, College Station, TX, USA
Keywords
fatty acids, canine, skin lipids, hair coat, total
fat, diet, linoleic acid, alpha-linolenic acid
Correspondence
John E. Bauer, Department of Veterinary Small
Animal Medicine and Surgery and Faculty of
Nutrition, College of Veterinary Medicine and
Biomedical Sciences, Texas A&M University,
College Station, TX 77843-4474, USA. Tel: 979
845 2351; Fax: 979 845 6978;
E-mail: jbauer@cvm.tamu.edu
Presented at the 10th Congress European
Society of Veterinary Comparative Nutrition
October 5–7, 2006, Nantes, France.
Received: 24 August 2007;
accepted: 3 April 2008
First published online: 12 August 2008
Introduction
Skin and hair coat (SHC)-related problems have his-
torically been observed in animals with nutritional
deficiencies and are often used as an indicator of a
companion animal’s nutritional status (Hansen and
Wiese, 1931; Bauer, 1994). Because dietary essential
fatty acid (EFA) deficiency results in matted hair
coat and unkempt appearance (Burr and Burr,
1929), these nutrients appear to be important to skin
Journal of Animal Physiology and Animal Nutrition 93 (2009) 505–511 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Verlag GmbH
Summary
It is generally believed that diets containing increased amounts of poly-
unsaturated fatty acids (PUFA) result in improved canine skin and hair
coat (SHC). However, the extent to which dietary fat amount and type
play a role remains to be systematically investigated. The objective of
this study was to investigate the role of both increased dietary fat
amount and type on SHC assessments of dogs. Improvements of SHC
conditions were investigated after feeding three diets containing
increased total dietary fat (i.e. 13% total fat) for 12 weeks in relation to
a lower fat acclimation diet (i.e. 9% total fat). The higher fat diets varied
in polyunsaturated and saturated fat types and amounts but total fat
was kept constant. Skin and hair coat assessments were performed at
selected intervals by a trained group of veterinarians and graduate stu-
dents. In addition, hair lipids were fractionated by thin layer chromatog-
raphy after extraction of plucked hair samples. Significant
improvements were found in hair coat glossiness and softness in all dogs
fed the higher fat diets in relation to the acclimation diet. Improvements
as a result of fat type were also seen but only at 12 weeks. A parallel
finding was a marked increase in hair cholesteryl ester content deter-
mined at the end of the study at which time SHC scores were signifi-
cantly improved. Skin and hair coat condition improvements may thus
be related to increased cholesteryl ester deposited on the hair shaft sur-
face when high fat diets are fed. Whereas this finding is preliminary,
hair lipid analysis may be a useful, non-invasive technique with which
to help assess dietary effects on canine SHC.
health. Providing dietary EFA assures epidermal
membrane fluidity, helps maintain the cutaneous
water permeability barrier, and supplies fatty acid
precursors of eicosanoids and other mediators of
normal cell function [National Research Council
(U.S.) Subcommittee on Dog Nutrition., 2006]. In
normal healthy skin, ceramides containing linoleic
acid (LA) are extruded as intercellular lamellar gran-
ules from epidermal keratinocytes which enhance
cell cohesion and impart an effective water barrier to
505
Role of increased dietary fat amount and type on SHC assessments of dogs
the epidermis The importance of LA to the epidermal
water barrier function was demonstrated in essential
fatty acid deficient rats and pigs in which depletion
of LA in ceramide phospholipid cellular fractions
directly correlated with increased epidermal water
loss (Elias et al., 1980; Wertz and Downing, 1982).
As LA is directly involved, many instances of dry,
dull hair coats, scaly, non-pruritic skin disorders
in dogs generally respond to dietary vegetable oil
supplements rich in this fatty acid (White, 1995).
In dogs, both serum and cutaneous fatty acid com-
position can be modified by dietary supplementation
(Campbell et al., 1992, 1995; Vaughn et al., 1994;
Campbell and Roudebush, 1995). Another dog study
compared whole ground linseed and sunflower seed
supplements in clinically normal dogs fed a basal
diet. In this case, significant short-term SHC score
improvements were found in both groups and both
supplements led to increased amounts of circulating
LA (Rees et al., 2001). Because plasma LA content
was greater in the linseed group, these findings sug-
gested that dietary ALA may exert a sparing effect
on LA conversion into other lipid metabolites
thereby allowing further LA accumulation.
Additional support for the importance of LA in
skin health was obtained when serum fatty acids
from atopic dogs were found to be significantly
reduced in LA in relation to normal animals (Seavik
et al., 2002). In these instances, impaired absorption
of LA may have existed as a result of decreased
serum triglyceride concentrations seen in atopic ani-
mals (Van denBroek and Simpson, 1990). Another
study found that cutaneous fatty acids in dogs with
seborrhea also have decreased LA content (Campbell
et al., 1992). Finally, other data suggest that an
interaction between zinc and LA may also exist
resulting in significantly enhanced canine SHC (Wat-
son, 1998; Marsh et al., 2000).
Hair coat sheen, glossiness and softness may be
influenced by the type and distribution of lipids pro-
duced by sebaceous glands. The major surface epi-
dermal lipids in dogs are cholesterol, cholesteryl
esters and wax diesters (Dunstan et al., 2000). Skin
surface lipids of dogs also contain a high proportion
of diol diesters which have lower mobility on thin
layer chromatography than wax diesters of other
species in spite of their similar fatty acid and diol
components (Sharaf et al., 1997). These lipids
become distributed over the hair shaft during hair
growth. In this way, glandular secretory activity may
potentially affect hair coat quality.
The objective of this study was to determine the
effects of increased total dietary fat containing
506 Journal of Animal Physiology and Animal Nutrition. ª 2008 The Authors. Journal compilation ª 2008 Blackwell Verlag GmbH
N. A. Kirby et al.
increased dietary EFA, including LA and ALA on
SHC condition scores in dogs. It was also hypothe-
sized that increased dietary fat type and amount
leads to sebum lipid modifications found on hair
lipid extracts.
Materials and methods
Animals and diets
Twenty-four clinically normal dogs, nine female bea-
gles and 15 male hound type mixed breeds, were
randomly assigned to one of three diet groups. There
were three beagles and five mixed breed dogs in
each group. The dogs were sexually intact and ran-
ged in age from 1.5 to 6.5 years with a median age
of 4 years. Body weights ranged from 15 to 40 kg
and body condition scores (BCS) were 3.1 ± 0.5
(SD) on a 5-point scale. Before initiation of the
study, all dogs were evaluated via complete blood
counts, serum biochemistry profiles, and serum thy-
roid stimulating hormone and free thyroxine con-
centration determinations. Laboratory tests showed
all dogs to be healthy and none were in an obese
body condition.
The dogs had initially been fed a commercial dry
extruded-type diet containing 14.5% fat (as-is) prior
to transferring them to another dry-extruded diet
containing 9% fat (as-is) for a 12-week acclimation
period (weeks )12 to 0). Amounts fed were based
on daily metabolic energy estimates calculated by
raising the body weight (kg) to the 0.75 power and
multiplying by the factor 132 [National Research
Council (U.S.) Subcommittee on Dog Nutrition.,
2006]. For the following 12 weeks (weeks 0–12), the
dogs were similarly fed one of three test diets (A, B
and C) containing approximately 13% fat (as-is). All
test diets were complete and balanced and of the
dry-extruded type and readily consumed. Body
weights and BCS were determined weekly and the
amounts fed were modified, where necessary, to
maintain BCS of approximately 3 out of 5 during
the feeding periods.
Diet A contained adequate amounts of EFAs (3.1 g
LA/1000 kcal, 0.42 g ALA/1000 kcal) and dietary
zinc (120 mg/kg). Diets B and C contained increased
but differing amounts of EFAs (Diet B, 9.3 g LA/
1000 kcal, 0.42 g ALA/1000 kcal); Diet C, 9.3 g LA/
1000 kcal, 3.3 g ALA/1000 kcal) and increased zinc
(both Diets B and C, 350 mg/kg). Total dietary fat in
all diets was approximately 13% (as-is). The diets
were isocaloric at approximately 3550 kcal/kg and
met the Association of American Feed Control Offi-
cials (AAFCO) standards for all nutrients (Table 1).
N. A. Kirby et al.
Table 1 Nutrient profile of the diets*
Nutrient Acclimation diet Diet A Diet B
Moisture (%) 12.0 6.2 6.6
Crude protein (%) 21.0 22.9 22.3
Crude fat (%) 9.0 13.1 12.9
Ash (%) 8.2 8.8 8.6
Crude fibre (%) 4.0 2.1 2.1
NFE (%) 45.8 46.9 47.5
ME (kcal/kg) 3250 3579 3565
NFE, nitrogen free extract; ME, metabolizable energy (calculated
according to Association of American Feed Control Officials (AAFCO),
2003).
*Values are % as-is basis except metabolizable energy (ME, kcal/
100 g). Ingredients for diets A, B and C included all of the following
(in order): lamb meal, ground rice, rice flour, rice bran, fat source (diet
A, beef tallow preserved with mixed tocopherols; diet B, sunflower oil
plus poultry fat, both preserved with mixed tocopherols; diet C,
ground flax seed plus poultry fat preserved with mixed tocopherols),
natural flavor, rice gluten, dried egg product, dried beet pulp, KCl,
L-lysine, dried kelp, NaCl, choline chloride, zinc sulphate, vitamin E
supplement, taurine, ferrous sulphate, ascorbic acid, biotin, copper
proteinate, niacin, manganous oxide, calcium pantothenate, vitamin
B12 supplement, riboflavin supplement, vitamin A, glucosamine hydro-
chloride, chondrotin sulphate, pyridoxine hydrochloride, thiamin mon-
nitrate, vitamin D3 supplement, menadione sodium bisulfite, calcium
iodate, folic acid.
Skin and hair coat condition scoring was per-
formed by a trained panel of seven evaluators using
a technique modified and expanded from that of
Rees et al. (2001). In the earlier study, reflectivity
and softness were combined into a single hair coat
score whereas in this work this assessment was
divided into two separate categories; glossiness and
softness. The greasiness category was retained but
additional assessments of scaliness and an overall
rating were added. Thus, scores for glossiness, greasi-
ness, scale, softness and overall coat quality were
obtained. An ordinal scoring system was used with a
1–5 scale (5, best in each category) assigning integer
values for each parameter. During the 12-week accli-
mation period, evaluations were performed at weeks
)12, )8, )4 and 0, whereas when the test diets were
fed, scoring occurred at weeks 0, 2, 4, 7 and 12.
Evaluators were blinded as to diet treatments used
after dogs were assigned to their respective test diets
groups.
Hair samples were collected for lipid analysis dur-
ing the test diet period at weeks 1, 3, 7 and 11 from
the proximal thigh region of the dogs. Hair samples
were collected by plucking hair with a haemostat.
To obtain a sufficient hair sample size, hair was
plucked from each dog from four to six times. The
hair samples were weighed and stored in sealed
Journal of Animal Physiology and Animal Nutrition. ª 2008 The Authors. Journal compilation ª 2008 Blackwell Verlag GmbH
Role of increased dietary fat amount and type on SHC assessments of dogs
tubes with teflon-lined caps with 2:1 (v/v) chloro-
Diet C
form/methanol containing 0.1% glacial acetic acid.
The samples were purged with N2 gas prior to cap-
6.2 ping the tubes and then stored at )20 C for 3–
22.9 4 weeks prior to lipid extraction. Hair from dogs of
13.3
each breed within a diet group was pooled at each
8.8
2.7
time-point to obtain sufficient sample for lipid analy-
46.1 sis. This resulted in three beagle group samples and
3569 three hound group samples at each of the four time-
points for lipid extraction and analysis (n = 6 per
time period).
Lipid analysis
Total lipid extractions of the canine hair samples
were performed using a modified Folch et al. (1957)
procedure as previously described (Bauer and Ran-
sone, 1983). Most of the lipid classes of sebum can
be resolved adequately using thin layer chromatog-
raphy on silica gel 60 (EM Science, Gibbstown, NJ,
USA) coated glass plates for both quantitative analy-
sis and for preparative separation using the proce-
dure of Downing (1968) as described by Sharaf et al.
(1997). Briefly, three sequential developing solvents
were used and plates dried in a nitrogen gas flow
box between solvents. The lipid extracts were frac-
tionated using development with hexane (to 17 cm),
followed by benzene (to 17 cm) and finally, hex-
ane:ether:acetic acid (50:50:1) to 9 cm. Lipid classes
were identified via authentic standards included on
each plate. After the plates were dry, they were
dipped in a 10% CuSO4 + 85% H3PO4 solution and
charred at 220 C. Individual lipid classes were
detected as charred spots. Lipids were quantified via
densitometry and external standardization with stan-
dard curves constructed for free cholesterol, wax
diester and cholesteryl ester.
Statistical analyses
Skin and hair coat score data were first assessed to
determine if evaluators correlated with one another
by calculating a Pearson’s correlation coefficient.
Correlations were considered acceptable whenever
the Pearson’s correlation coefficient was ‡0.30 at
p < 0.05. Because this type of data is subjective and
based on individual opinion, this criteria is regarded
as acceptable, albeit weakly correlated, but suitable
in the behavioral sciences (Cohen et al. (2003).
Using these criteria, it was found that all evaluators
correlated with one another to proceed with further
statistical analyses. Kruskal–Wallis one-way, non-
parametric analysis of variance was used to test for
507
Role of increased dietary fat amount and type on SHC assessments of dogs
significant differences over all time-points during
both the acclimation and test periods and also
among the diets. Repeated measures anova was used
to evaluate the hair lipid data for main time, diet
and time*diet effects followed by Tukey’s multiple
comparison testing where appropriate.
Results
Amounts of food consumed were similar among the
animals fed the experimental diets in both the bea-
gles and hound-type dogs and all animals main-
tained constant body weights during the study (data
not shown). Body condition scores at the end of the
test diet period averaged 3.1 ± 0.5 (SD) and were
unchanged from those seen at the acclimation base-
line.
During the acclimation period, statistically signifi-
cant decrements were seen in all SHC assessment
categories except for scaliness which remained con-
stant throughout this period (Fig. 1). These differ-
508 Journal of Animal Physiology and Animal Nutrition. ª 2008 The Authors. Journal compilation ª 2008 Blackwell Verlag GmbH
N. A. Kirby et al.
ences reflected the change to the acclimation diet. It
should be noted that these scores stabilized between
4 weeks (week )8) and 8 weeks (week )4) indicat-
ing that all dogs had achieved a new steady state
during this time. When the test diets were fed, statis-
tical analyses revealed improvements in glossiness,
softness and overall assessment scores beginning at
week 4 which again stabilized at week 7. Significant
improvement in greasiness score did not occur until
week 7, however, and scaliness scores were
unchanged but quite variabile at weeks 0 and 2
(Fig. 1).
Although significant time effects were observed
throughout the test diet period, statistically signifi-
cant diet effects on SHC assessments were not seen
until week 12. Kruskal–Wallis one-way non-para-
metric analysis of variance performed at week 12
revealed statistically significant overall hair coat
improvement with the high n-6 fatty acid containing
diet (diet B) in relation to the high saturated fat diet
(diet A). Also, improved scaliness and softness scores
Fig. 1 Skin and hair coat scores (mean val-
ues) during the acclimation and test diet peri-
ods. Statistically significant time effects were
observed for all test diets for glossiness, soft-
ness and overall scores. Numbers not in com-
mon denote significant differences over time
during the diet acclimation period (weeks
)12 to 0), p < 0.05. During the test diet per-
iod, letters not in common denote statistically
significant differences over time (weeks 0–12)
p < 0.05. Sample size is 168 at each time-
point (seven evaluators · 24 dogs). Error bars
have been omitted for clarity. Overall stan-
dard deviations for each assessment were –
acclimation period: glossiness, 0.80; softness,
0.83; greasiness, 0.83, scaliness, 0.95 and
0.78, overall; test diet period: glossiness,
0.76; softness, 0.80; greasiness, 0.80, scali-
ness, 0.85 and 0.85, overall. No diet effects
were observed at weeks 0, 2, 4 or 7. See
Fig. 2 for significant diet effects observed at
week 12.
N. A. Kirby et al. Role of increased dietary fat amount and type on SHC assessments of dogs

Fig. 2 Skin and hair coat scores (mean ± SEM) at the end of the test
diet period (week 12). Letters not in common for a skin and hair coat
assessment category are significantly different at p < 0.05, n = 8 per
diet group.

were seen with the high n-6 plus n-3 diet (diet C)
in relation to diet A. No significant diet effects for
glossiness or greasiness were observed among the
test diets (Fig. 2).
Changes in the major lipid fractions from extracts
of hair samples were also observed, over time, with Fig. 3 Representative thin layer chromatograms of the hair lipid
the test diets. The major lipid classes identified in extracts from pooled samples in each diet group. Duplicate analyses
hair were cholesterol, wax diester (i.e. dog diester as are shown. The left lane of each chromatogram contains the following
standards: squalene; CE, cholesteryl ester; WE, wax ester; ME, methyl
described by Sharaf et al., 1997) and cholesteryl
ester; TG, triacylglycerol; FFA, free fatty acid; DG, diacylglycerol; MG,
esters (CE). Markedly increased CE contents were monoacylglycerol; PL, phospholipids. The position of the wax diester
visually apparent on the thin layer chromatograms as described by Sharaf et al. (1997) is indicated as Dog Diester. Note
with the test diets in both dog breeds studied (Fig. 3, the lack of CE fraction at week 1 of the test diet period in relation to
weeks 1 and 11 shown). Because diet effects were week 11.
not seen for the first 11 weeks of the test diet period,
lipid data from each of the two breeds within each
diet group were pooled for further analysis at each
time-point resulting in a sample size of 6 per group.
Repeated measures anova of the resultant data
revealed a statistically significant increase in hair CE
content over time (Fig. 4). In addition, the ratio of
CE to wax diesters (WD) was also significantly
increased primarily because of the increase in CE
content. It is noteworthy that these results were
seen at the same time that improvements of SHC
scores were observed. However, lack of larger sample
sizes prevented any further correlation analyses to
be performed.

Discussion
During the acclimation period, statistically significant
decrements in glossiness, greasiness, softness and
overall assessment scores were seen at approximately
4 weeks, remaining relatively constant thereafter.
Fig. 4 Concentrations of hair lipid extract fractions and CE/WD ratio
during the test diet period. Letters not in common for a given fraction
are significantly different, p < 0.05. CE, cholesteryl ester; CH, free cho-
lesterol; WD, wax diester. Sample size is 6 for each time-point.

Journal of Animal Physiology and Animal Nutrition. ª 2008 The Authors. Journal compilation ª 2008 Blackwell Verlag GmbH 509
Role of increased dietary fat amount and type on SHC assessments of dogs
During the test period, significant increases in glossi-
ness, softness and overall scores were subsequently
seen, again beginning at 4 weeks, remaining
unchanged until the 7-week assessment, then
improving further at 12 weeks. Significant improve-
ments in greasiness scores were not observed until
7 weeks. In an earlier study, Campbell et al. (1995)
studied fatty acid profiles of plasma and skin after
dietary fat modification and concluded that 12 weeks
was necessary for skin lipids to reflect dietary altera-
tions. However, in that study, skin samples were
analysed only at 6 and 12 weeks and no data were
obtained between these two time periods. Another
study fed dogs varying dietary polyunsaturated fat
types for 8 weeks before assessing SHC and found
increased hair coat sheen and decreased tran-epider-
mal water loss (Campbell and Roudebush, 1995). By
contrast, Rees et al., 2001 reported that SHC
improvements occurred at 4 weeks after dietary fat
modification as has been found in the present inves-
tigation. However, in this study, the initial signifi-
cant changes which appeared as early as 4 weeks
remained so at 7 weeks. Thus, in a veterinary prac-
tice setting, it may be prudent to recommend a per-
iod of 6–8 weeks after dietary fat modification before
SHC responses are evaluated to fully ascertain the
effects of such a dietary change.
All test diets contained higher total fat in relation
to the acclimation diet. Thus, the initial improve-
ments seen, over time, in glossiness, softness and
overall coat quality appear to be because, at least
in part, of these higher total dietary fat concentra-
tions. Separate diet effects were also observed, but
only at 12 weeks, in which statistically significant
differences because of polyunsaturated fat type
occurred. Overall hair coat improvement was found
with the high n-6 fatty acid containing diet (diet
B) and improved scaliness and softness scores seen
with the high n-6 plus n-3 diet (diet C) in relation
to diet A which contained high saturated fat. Con-
sequently, while increased total dietary fat was ini-
tially found to improve most of the canine SHC
scores performed in this study, additional incre-
mental improvements were observed when the
higher fat diets also included relatively more poly-
unsaturated fats. Because numerous additional ben-
efits of including polyunsaturated fatty acids in
canine diets exist including eicosanoid production
and neurologic function (reviewed in Bauer, 2006),
the use of properly balanced combinations of all
fatty acid types including n-6 and n-3 fatty acids
along with higher total dietary fat may be preferred
overall.
510 Journal of Animal Physiology and Animal Nutrition. ª 2008 The Authors. Journal compilation ª 2008 Blackwell Verlag GmbH
N. A. Kirby et al.
Of additional interest in this study were changes
in the major lipid fractions of hair extracts during
the test diet period. Statistically significant increases
in hair total cholesteryl ester (CE) concentrations
and the CE/WD ratio were found in all three test
diet groups at the end of the feeding period. This
ratio may be a useful way in which to normalize
data in future studies where external standardization
is not employed. Whether these lipid alterations cor-
relate with improvement of SHC assessments is
unknown at this time because hair lipid extracts
were pooled and sample size was limited. Conse-
quently, correlation data are not available. Nonethe-
less, these initial data show increased hair surface
lipid CE in parallel with improvements in SHC
assessment scores when higher fat diets were fed.
Should a positive correlation exist between these
two parameters, hair lipid analysis may provide a
useful, non-invasive technique to quantify dietary
effects on SHC. An earlier study also demonstrated
increased sebum CE fraction in dogs fed diets con-
taining increased total fat (Dunstan et al., 2000).
Furthermore, we have shown that increased total
dietary fat also elevates plasma CE in dogs (Bauer
et al., 1997; McAlister et al., 1996). These findings
support the possibility that elevation of CE of hair
surface lipids may be a useful objective marker of
the more subjective assessments of hair coat glossi-
ness, softness and overall condition.
Acknowledgements
The authors acknowledge the technical assistance of
Karen Bigley and Angela Wright-Rodgers during this
investigation.
References
Association of American Feed Control Officials (AAFCO).
2003: Official Publication. AAFCO Official Publication,
Atlanta, GA.
Bauer, J. E., 1994: The potential for dietary polyunsatu-
rated fatty acids in domestic animals. Australian Veteri-
nary Journal 71, 342–345.
Bauer, J. E., 2006: Facilitative and functional fats in diets
of dogs and cats. Journal of the American Veternary Medi-
cal Association 229, 680–684.
Bauer, J. E.; Ransone, W. D., 1983: Fatty acid composi-
tion of serum lipids in fasting ponies. Lipids 18, 397–
401.
Bauer, J. E.; McAlister, K. G.; Rawlings, J. M.; Markwell,
P., 1997: Molecular species of cholesteryl esters formed
N. A. Kirby et al.
via plasma lecithin-cholesterol acyltransferase in fish
oil supplemented dogs. Nutrition Research 17, 861–872.
Burr, G.; Burr, M. M., 1929: A new deficiency disease
produced by the rigid exclusion of fat from the diet.
Journal of Biological Chemistry 82, 345–367.
Campbell, K. L.; Roudebush, P., 1995: Effects of four
diets on serum and cutaneous fatty acids, transepider-
mal water losses, skin surface lipids, hydration and
condition of the skin and haircoat of dogs. In: Proceed-
ings, Annual AAVD & ACVD Meeting, Santa Fe, NM.
pp. 80–81.
Campbell, K. L.; Uhland, C. F.; Dorn, G. P., 1992: Effects
of oral sunflower oil on serum and cutaneous fatty acid
concentration profiles in seborrheic dogs. Veterinary
Dermatology 3, 29–35.
Campbell, K. L.; Czarnecki-Maulden, G. L.; Schaeffer, D.
J., 1995: Effects of animal and soy fats and proteins in
the diet on fatty acid concentration in the serum and
skin of dogs. American Journal of Veterinary Research 56,
1456–1459.
Cohen, J.; Cohen, P.; West, S. G.; Aiken, L. S., 2003:
Applied Multiple Regression/Correlation Analysis for the
Behavioral Sciences, 3rd edn. Lawrence Erlbaum
Associates, Hillsdale, NJ.
Downing, D. T., 1968: Photodensitometry in the thin-
layer chromatographic analysis of neutral lipids. Jour-
nal of Chromatography 38, 91–99.
Dunstan, R. W.; Herdt, T. H.; Meu, L.; Credille, K. M.;
Kennis, R. A.; Maier, R. L.; Dziezyc, J.; Tucker, K. A.;
Reinhart, G. A.; Davenport, G. M., 2000: The role of
nutrition on canine sebum secretion: a preliminary
report. In: G. A. Reinhart, D. P. Carey (eds), Recent
Advances in Canine and Feline Nutrition, Vol 3. Iams
Nutrition Symposium, Orange Frazer Press, Wilming-
ton, OH, pp. 23–35.
Elias, P. M.; Brown, B. E.; Ziboh, V. A., 1980: The perme-
ability barrier in essential fatty acid deficiency: evidence
for a direct role for linoleic acid in barrier function. Jour-
nal of Investigative Dermatology 81, 230–233.
Folch, J.; Lees, M.; Stanley, G. H. S., 1957: A simple
method for the isolation and purification of total lipids
from animal tissues. Journal of Biological Chemistry 226,
497–509.
Hansen, A. E.; Wiese, H. F., 1931: Fat in the diet in rela-
tion to nutrition of the dog. I. Characteristic appear-
ance and changes of animals fed diets with and
without fat. Texas Reports in Biology and Medicine 9,
491–515.
Journal of Animal Physiology and Animal Nutrition. ª 2008 The Authors. Journal compilation ª 2008 Blackwell Verlag GmbH
Role of increased dietary fat amount and type on SHC assessments of dogs
Marsh, K. A.; Ruedisueli, F. L.; Coe, S. L.; Watson, T. D.
G., 2000: Effects of zinc and linoleic acid supplementa-
tion on the skin and coat quality of dogs receiving a
complete and balanced diet. Veterinary Dermatology 11,
277–284.
McAlister, K. G.; Bauer, J. E.; Harte, J.; Rawlings, J.;
Markwell, P., 1996: Canine plasma lipoproteins and
lecithin-cholesterol acyltransferase activities in dietary
oil supplemented dogs. Veterinary Clinical Nutrition 3,
50–56.
National Research Council (U.S.) Subcommittee on Dog
Nutrition, 2006: Fat and fatty acids. In: Committee on
Animal Nutrition (eds), Nutrient Requirements of Dogs.
National Academies Press, Washington, DC,
pp. 81–110.
Rees, C. A.; Bauer, J. E.; Burkholder, W. J.; Kennis, R.
A.; Dunbar, B. L.; Bgley, K. E., 2001: Effects of dietary
flax seed and sunflower seed supplementation on nor-
mal canine serum polyunsaturated fatty acids and skin
and hair coat condition scores. Veterinary Dermatology
12, 111–117.
Seavik, B. K.; Thossesen, S. I.; Taugbol, O., 2002: Fatty
acid composition of serum lipids in atopic and healthy
dogs. Research in Veterinary Science 73, 153–158.
Sharaf, D. M.; Clark, S. J.; Downing, D. T., 1997: Skin
surface lipids of the dog. Lipids 12, 786–790.
Van denBroek, A. H. M.; Simpson, J. W., 1990: Fat
absorption in dogs with atopic dermatitis. In: C. von
Tscharner, R. E. W. Halliwell (eds), Advances in Veteri-
nary Dermatology, Vol 1. Bailliere Tindall, London,
pp. 155–160.
Vaughn, D. M.; Reinhart, G. A.; Swaim, S. F.; Lauten, S.
D.; Garner, C. A.; Boudreaux, M. K.; Spano, J. S.; Hoff-
man, C. E.; Conner, B., 1994: Evaluation of effects of
dietary n-6 to n-3 fatty acid ratios on leukotriene B
synthesis in dog skin and neutrophils. Veterinary Derma-
tology 5, 163–173.
Watson, T. D. G., 1998: Diet and skin disease in dogs and
cats. Journal of Nutrition 128, 2783S–2789S.
Wertz, P. W.; Downing, D. T., 1982: Glycolipids in mam-
malian epidermis: structure and function of the water
barrier. Science 217, 1261–1262.
White, P., 1995: Essential Fatty Acids in Veterinary Medicine.
Monograph, Bayer Corporation, Agricultural Division,
Animal Health Veterinary Learning Systems, Shawnee
Mission, KS, 1–48.
511