The

Veterinary Journal
The Veterinary Journal 172 (2006) 78–85
www.elsevier.com/locate/tvjl

A quantitative study of the equine soft palate

using histomorphometry
Lucy E. Richardson 1, Glenn K. Wakley, Samantha H. Franklin *

Department of Anatomy, University of Bristol, School of Veterinary Science, Southwell Street, Bristol BS2 9EJ, UK

Accepted 16 February 2005

Abstract

Dorsal displacement of the soft palate is a common cause of upper airway obstruction in racehorses and is of unknown aetiology.
To determine whether the palate may displace for structural reasons, knowledge of the normal soft palate is required. The present
study aimed to describe, qualitatively and quantitatively, the structure of the normal equine soft palate using histomorphometry. In
soft palates from 12 Thoroughbreds, glandular tissue predominated (ca. 40% of total area), located mainly in the rostral and ventral
regions. Rostrally, muscles attached to a tendinous aponeurosis located dorsal to the glandular tissue. Muscle was most abundant in
the dorsal mid region and decreased caudally. The oral mucosa consisted of a non-keratinised stratified squamous epithelium
whereas the nasopharyngeal mucosa was pseudostratified, columnar and ciliated. Elastin fibres were observed in the nasopharyngeal
submucosa, becoming more prevalent caudally. The palates were bilaterally symmetrical although the proportion of tissue types
varied considerably between individuals.
 2005 Elsevier Ltd. All rights reserved.

Keywords: Soft palate; Histomorphometry; Anatomy; Image analysis; Horse

1. Introduction This leads to a reduction in ventilation and gas ex-

change, and is associated with impaired athletic perfor-
The soft palate acts to separate the nasopharynx from mance (Franklin et al., 2002).
the oropharynx and plays an important role in both Dorsal displacement of the soft palate (DDSP) is now
breathing and deglutition. In the horse, a unique rela- recognised to be one of the most common upper respira-
tionship exists between the larynx and soft palate. The tory tract disorders affecting the racehorse (Beard, 1996;
caudal border of the soft palate lies ventral to the rostral Martin et al., 2000). Although a number of theories have
body of the epiglottis, except during deglutition, render- been proposed as to the cause of DDSP, including phys-
ing this species an obligate nasal breather (Negus, 1949). ical, behavioural and pathological abnormalities, its
In some horses, however, the soft palate may become underlying aetiology remains unclear. Recently, the con-
displaced dorsal to the epiglottis during exercise, causing dition has been likened to that of obstructive sleep ap-
an obstruction to airflow (Morris and Seeherman, 1990). noea (OSA) in humans (Holcombe et al., 1998).
Study of the normal structure of the soft palate al-
lows identification or elimination of structural factors
*
Corresponding author. Tel.: +44 117 928 9361; fax: +44 117 928 which may have a bearing on pathological mechanisms.
9622. This has been addressed in the studies of the human soft
E-mail address: [email protected] (S.H. Franklin).
1
Present address: Department of Veterinary Clinical Sciences, The
palates where alterations in pharyngeal structure and
Royal Veterinary College, Hawkshead Lane, North Mymms, Hatfield, function are considered fundamental in the pathogenesis
Hertfordshire AL9 7TA, UK. of snoring and OSA (Kuehn and Kahane, 1990; Ettema

1090-0233/$ - see front matter  2005 Elsevier Ltd. All rights reserved.
doi:10.1016/j.tvjl.2005.02.032
 L.E. Richardson et al. / The Veterinary Journal 172 (2006) 78–85 79

and Kuehn, 1994; Metternich et al., 2000). Similarly for saline for 24 h, to account for shrinkage. After fixation,
investigation of DDSP, a sound knowledge of the anat- the soft palates were bisected down the midline. The
omy of the normal soft palate is essential to the under- right and left halves were divided into 10 equal sized tis-
standing of the underlying aetiology. Although the sue blocks from rostral to caudal with the face of the
gross anatomy of the equine soft palate and its sur- block being perpendicular to the oral and nasal surfaces.
rounding structures has been documented qualitatively Each block was the thickness of 10% of the length of the
(Sisson, 1975; Cook, 1981) little attention had been paid whole soft palate (Fig. 1). This allowed for normalisa-
to its detailed or quantitative histological examination, tion of soft palate lengths between subjects.
yet it is acknowledged that the detailed morphology of In five cases, both left and right sides were examined
the velopharyngeal region is best studied using histolog- in order to assess bilateral symmetry. Having established
ical techniques (Kuehn and Kahane, 1990). that there were no significant differences between tissues
The aim of this study was to characterise qualitatively of the left and right sides, only selected blocks were eval-
and quantitatively the tissues of the normal equine uated for the remaining horses. In one subject, the sec-
soft palate by means of light microscopy and ond side was sectioned longitudinally to enable
histomorphometry. qualitative assessment of tissue changes between and
within the transverse tissue blocks.
Individual tissue blocks were embedded in paraffin
2. Materials and methods and 5 lm sections were cut from the intervening face
of each block. These were stained with haematoxylin
The heads of 12 Thoroughbred horse cadavers were and eosin (H and E) and MassonÕs trichrome to assess
obtained from an abattoir. The soft palate from each general tissue architecture, periodic acid-Schiff (PAS)
animal was dissected out in its entirety from the oral as- and alcian blue to assess mucin type within the glandu-
pect. Care was taken to maintain pharyngeal tissue lat- lar tissue, and VerhoeffÕs and Van Gieson stains for
erally and resect as close to the hard palate as possible assessment of elastin fibres.
rostrally. The excised palates were examined to confirm The slides were assessed qualitatively and quantita-
that there were no signs of disease or previous surgery. tively using a Sony 3 CCD colour video camera
They were then prepared using a method similar to that mounted on a Leica DM/RB light microscope and the
described for human palatal tissue (Kuehn and Kahane, image displayed on a computer monitor. For qualitative
1990). assessment, the slides were examined using a high mag-
The soft palates were measured along the midline, be- nification (40·) to enable determination of detailed
fore and after fixation in 10% neutral buffered formol structures (such as epithelia and elastin fibres), and with

Fig. 1. Schematic diagrams of the plan for histological sectioning. Transverse sections were taken at 10% increments along the length of the soft
palate. (L1–L5 are the longitudinal sections).
80 L.E. Richardson et al. / The Veterinary Journal 172 (2006) 78–85

medium magnifications (20· and 10·) for analysing Glandular tissue symmetry in the equine soft palate
other tissues. 90

Mean % ti ssue present

80
For the quantitative measurements a low magnifica- 70
tion (5·) was used. Three columns of fields were exam- 60
50
ined from the respiratory surface to the oral surface. 40
One column was taken at the most medial edge and 30
20
one at the most lateral edge of the tissue section. A third 10
column was measured mid-way between these. The dif- 0
ferent tissue types measured included: glandular and 10% 20% 30% 40% 50% 60% 70% 80% 90%

lymphoid tissue, muscular tissue, connective tissue (a) % Length of the soft palate from the hard palate

(including the tendon aponeurosis) and other tissue (epi-

Muscle tissue symmetry in the equine soft palate
thelia and adipose tissue remnants). The tissue types in 60
each column were recorded and quantitative measure-

Mean % tissue present

50
ments of the section image were achieved by moving a 40
cursor over a digitising bit-pad (CalComp, Osteometrics
30
Inc.). This produced a line tracing around the tissue
20
chosen on the computer screen. The line length and area
10
enclosed by the line was then calculated using Osteo-
0
measure software (Osteometrics Inc.). 10% 20% 30% 40% 50% 60% 70% 80% 90%

(b) % Length of soft palate from the hard palate

3. Data analysis Fig. 2. Comparison of glandular (a) and muscular (b) tissue of left
(white) and right (black) side soft of the soft palate. (values are
expressed as mean ± sd; 10% is rostral and 90% is caudal).
The mean (±sd) percentage of each tissue type was
calculated for all 12 palates. The 12 palates were divided
into young (<10 years) or old (P10years) and by gen-
der, and differences in tissue types assessed using a and mucus glands. Lymphocytes were scattered in the
two-sample t test. Left and right symmetry was assessed respiratory mucosa and submucosa (Fig. 3a).
in five horses using a paired StudentÕs t test. Statistical The oral mucosa was non-keratinised stratified squa-
significance was set at P < 0.05. Intra-observer and mous epithelium that became more folded caudally. The
inter-observer error was assessed. The variability for this oral submucosa largely comprised connective tissue,
data was investigated using Pearson correlation; intra- which had finger-like projections into the epithelium.
observer error of all tissues was found to have an aver- Some glandular tissue was present and lymphoid tissue
age r-value of 0.98 and the inter-observer r-value was was found aggregated around mucosal crypts (tonsilla
0.87. veli palatini) (Fig. 3b).
Elastin fibres were found mostly in the nasopharyn-
geal submucosa and surrounding muscle bundles within
4. Results the soft palate. These were graded according to a semi-
quantitative scale (Table 1, Fig. 4). The fibres were
The specimens included soft palates from seven males found to increase in both prevalence and thickness in
and five females, with a mean age of 10.7 years (±5.7 sd). the caudal regions with fewer, finer fibres present in
The mean length of the soft palate was 134 mm the rostral sections (Fig. 5).
(±12 mm sd) before fixation and 124 mm (±8 mm) after Glandular tissue was predominant in the rostral soft
fixation. No significant differences were found in soft palate (up to 40% of the length of the soft palate from
palate length when they were compared by gender the hard palate) and decreased caudally (Figs. 2,6 and
(P = 0.17) or age (P = 0.23). 7). The glands were found to be predominantly mucous
The soft palates from five subjects were assessed for glands with a tubular and tubulo-alveolar structure, and
bilateral symmetry. The results for muscular and glan- divided into lobules by connective tissue septa. Interlob-
dular tissue are shown in Fig. 2. There was no significant ular and excretory ducts were also evident. PAS/alcian
difference for any of the tissues between the left and blue staining revealed that acidic mucins were present
right sides. only (Fig. 8).
Qualitative examination revealed that the nasopha- A tendinous aponeurosis (Fig. 7a and Fig. 9) was
ryngeal mucosa was composed of pseudostratified present in the rostral sections, up to 40% of the length
columnar ciliated (respiratory) epithelium. This was of the soft palate from its attachment to the hard palate,
highly folded and thinner than the oral mucosa. The although the exact length varied between individuals.
submucosa consisted of connective and lymphoid tissues Lateral fibres of the m. tensor veli palatini tendinous
 L.E. Richardson et al. / The Veterinary Journal 172 (2006) 78–85 81

Fig. 3. (a) The columnar ciliated epithelium of the nasal mucosa. Scattered lymphocytes can be seen (arrows). MassonÕs Trichrome stain.
Magnification 40· (scale bar = 0.05 mm). (b) The oral mucosa (OM) of the rostral soft palate containing lymphoid tissue (arrows). MassonÕs
Trichrome stain. Magnification 25· (scale bar = 1 mm).

Table 1 insertion were seen at the caudolateral edge of the

Subjective scale for analysis of elastin fibres aponeurosis.
Grade Description Muscular tissue was predominant in the midsections
1 Few sparse fine elastin fibres present, interspersed widely (Figs. 2, 6 and 7). Longitudinal fibres of the palatinus
throughout the connective tissue muscle originated from the aponeurosis and extended
2 Elastin fibres more densely packed in discrete bundles, but along the length of the soft palate, in the dorsal midline.
bundles are widely dispersed throughout the connective
The m. palatinus was surrounded by a connective tissue
tissue
3 Elastin fibres in densely packed bundles but found widely sheath onto which some of the other palatal muscles in-
interspaced in most of the connective tissue serted. Lateral muscle fibres of the m. levator veli pala-
4 Many densely packed elastin fibre bundles found close tini formed a thin but broad insertion in the midline,
together throughout most of the connective tissue. Fibres ventral to the palatinus muscle, in the region between
are now much thicker
50% and 80% of the soft palate length; the left and right

Fig. 4. VerhoeffÕs and Van Gieson stain showing the elastin fibres (stained black) within the soft palate. Connective tissue is shown in red. Objective is
40·. Scale bar = 0.05 mm.
82 L.E. Richardson et al. / The Veterinary Journal 172 (2006) 78–85

Elastin fibre grading in the equine soft palate ture comparisons of soft palates from horses with
5 abnormal palatal function.
In five subjects the percentages of the different tissue
Elastin gnade

4
types were compared between left and right sides. There
3
was no significant difference between the two sides for
2 any tissues in any of the subjects, suggesting that tissues
1
within the normal equine soft palate are symmetrical.
Considerable inter-subject variation was observed in
0 the proportion of tissue types within the soft palate, as
10% 20% 30% 40% 50% 60% 70% 80% 90%
indicated by the fairly large standard deviations. How-
% Length of the soft palate from the hard palate
ever, no obvious differences were observed between
Fig. 5. Mean ± sd elastin fibre grading along the length of the soft horses of different gender or age.
palate (10% is rostral and 90% is caudal). The predominant tissue in the equine soft palate was
glandular. The structure of the glandular tissue, which
consisted of predominantly mucous glands that con-
tained only acidic mucins, was similar to that described
Other Tissue Muscle in other species (Black, 1977; Finkelstein et al., 1992).
Connective Tissue Glandular Tissue Glandular tissue was most predominant within the ros-
tral soft palate and decreased towards the caudal bor-
Tissue distribution in the soft palate
der. This pattern differs from that observed in the
100% human soft palates, where glandular tissue averages
Mean % tissue present

about 22% of the total tissue volume, a proportion that

80% remains fairly constant along the whole length of the
structure (Kuehn and Kahane, 1990; Ettema and
60% Kuehn, 1994). In the present study, the glandular vol-
40%
ume ranged from 70% rostrally to 20% caudally and
the average glandular volume across the whole palate
20% was approximately 40%, almost twice that of humans.
This difference may have an underlying functional ba-
0% sis since salivary gland size can undergo selective hyper-
10% 20% 30% 40% 50% 60% 70% 80% 90% trophy or atrophy depending on functional stimulation
% Length of soft palate from the hard palate (Wells and Peronace, 1967; Hall and Schneyer, 1977).
In rats, a dry diet consisting primarily of cellulose led
Fig. 6. Total tissue composition along the length of the soft palate
(average of 12 subjects).
to an increase in salivary gland weight compared with
that in rats fed a liquid diet where there was cessation
of gland growth (Wells and Peronace, 1967). In the
sides forming a sling-like structure beneath the palatinus horse the diet consists primarily of large amounts of
muscle. The m. palatopharyngeus was observed to arise dry roughage which could lead to a functional need
from the aponeurosis lateral to the m. palatinus and fur- for increased saliva production and therefore a large
ther fibres were found to attach ventrally onto the amount of glandular tissue. Further research is required
sheath of the m. palatinus, along its length. to determine whether there are differences between
The proportions of each tissue type varied between horses fed different diets.
individuals (Table 2). However, no significant gender- The distribution of muscular tissue within the soft
or age-related changes were found for any tissue type. palate is of great interest with respect to its biomechan-
ical properties. A number of intrinsic muscles are recog-
nised within the soft palate; the co-ordinated
5. Discussion contraction of which are important in determining its
position both during breathing and deglutition.
The aim of this study was to provide qualitative and The proportion of muscular tissue was found to vary
quantitative information on the structure of the equine along the length of the soft palate. This was similar to
soft palate. The methodology involved gross dissection the finding in human soft palate (Ettema and Kuehn,
and assessment as well as microscopic and morphologi- 1994). No muscle was observed rostrally. The palatal
cal analysis of tissues within the soft palate. The preci- muscles arose from a broad tendinous aponeurosis ros-
sion of the histomorphometry technique was high, as trally and the bulk of the musculature was located dor-
determined by the inter- and intra-observer reliability, sally in the mid regions of the soft palate, and again
which indicates that this may be a useful method for fu- diminished caudally.
 L.E. Richardson et al. / The Veterinary Journal 172 (2006) 78–85 83

Fig. 7. Sections along the length of the soft palate (SP), stained with MassonÕs Trichrome. The nasal surface is at the top of each photograph. Scale
bar = 1 cm.

Fig. 8. Glandular tissue stained with PAS/Alcian Blue shows mucus Fig. 9. Tendinous aponeurosis stained with MassonÕs Trichrome.
glands with acid mucins present. Magnification 10· (scale Magnification 40· (scale bar = 0.05 mm).
bar = 0.25 mm).

The palatinus muscle is a paired muscle lying in the induce instability of the rostral portion of the soft palate
dorsal midline. It was enclosed within a connective tis- in exercising horses (Holcombe et al., 1997).
sue sheath along its length, and it has been suggested The palatopharyngeus muscle also provides lateral
that this feature may constitute the intrinsic support of support to the soft palate. Contraction of this muscle,
the dorsal aspect of the soft palate, controlling its stiff- together with the contraction of palatinus muscle, re-
ness and preventing distortion (Kuehn et al., 1988). sults in shortening of the soft palate and depression of
Fibres of the tensor veli palatini muscle inserted later- the caudal portion towards the tongue (Kuehn et al.,
ally onto the caudal aspect of the aponeurosis. The ac- 1982). Fibres of the palatopharyngeus muscle arose ros-
tion of this muscle is to tense the rostral portion of the trally from the aponeurosis, lateral to the origin of the
soft palate and depress it towards the tongue, thereby palatinus muscle. Additional fibres were found along
opposing the suction effect within the nasopharynx dur- the length of the soft palate where they attached in the
ing inspiration (Heffron and Baker, 1979). Experimental midline, ventral to the levator veli palatini and the pal-
bilateral transection of the tendons has been shown to atinus muscles. This differs from the distribution of the
84 L.E. Richardson et al. / The Veterinary Journal 172 (2006) 78–85

Table 2
Mean (+sd) percentage tissue area of the different tissue types in each block along the length of the soft palate
Tissue block (%) Tissue type
Glandular Muscular Connective tissue Other
10 71.0 (6.7) 0 24.1 (6.5) 4.9 (3.4)
20 67.8 (7.7) 0 26.9 (7.0) 5.3 (2.2)
30 66.4 (7.5) 1.0 (1.38) 28.2 (5.6) 4.4 (1.6)
40 47.2 (6.9) 11.3 (11.5) 35.2 (4.2) 6.3 (4.0)
50 40.8 (8.6) 31.2 (8.3) 23.5 (4.8) 4.5 (2.2)
60 23.3 (11.7) 50.7 (13.2) 22.7 (6.3) 3.3 (1.7)
70 23.7 (8.9) 38.6 (11.3) 30.2 (10.1) 7.5 (4.8)
80 20.5 (5.7) 28.8 (6.8) 40.2 (4.8) 10.5 (2.4)
90 20.0 (5.6) 26.9 (6.2) 43.5 (5.9) 9.6 (1.6)

muscle in human palates where it does not extend as far content was found to increase in density and fibre thick-
caudally (Kuehn and Kahane, 1990). These differences ness towards the caudal border of the soft palate. It is
are likely to occur because of the fact that the equine proposed that the elastin fibres are probably necessary
soft palate is relatively longer than that of the human, to allow deformation of the caudal soft palate by the food
with the free border extending to the base of the epiglot- bolus during swallowing and may also have a role in the
tis (Sisson, 1975). It is this arrangement which renders maintenance of the patency of the nasopharyngeal air-
the horse an obligate nasal breather, compared with way by helping to prevent dorsal billowing of the soft pal-
other species, including humans which can switch from ate (Kuehn and Kahane, 1990; Ahern, 1994).
nasal to oral breathing (Negus, 1949). Structural differences have been reported in humans
Fibres of the levator veli palatini muscle occurred in with palatal dysfunction leading to snoring and OSA. Pa-
the region between 50% and 80% of the length of the tients with snoring have significantly greater proportions
soft palate and formed a broad insertion onto the palati- of fat and connective tissue in combination with muscle
nus sheath in the midline, ventral to the palatinus mus- atrophy (Metternich et al., 2000). Other studies have also
cle, forming a sling-like structure as described in humans reported mucous gland hypertrophy (Woodson et al.,
(Kuehn and Kahane, 1990; Moon et al., 1998). The 1991) and neurogenic lesions resulting in atrophy of the
function of the levator is to raise the soft palate during palatal musculature (Woodson et al., 1991; Friberg
deglutition and vocalisation and to aid oral ventilation et al., 1998). It is possible that similar differences may
in non-obligate nasal breathers (Kuehn et al., 1982). occur in horses with palatal dysfunction. Indeed, recent
Connective tissue accounted for a large proportion of reports from HolcombeÕs group suggest that some horses
the soft palate, ranging from 20% to 40% along its with intermittent DDSP may have altered EMG activity
length. The nasopharyngeal mucosa consisted of pseud- of the palatinus and palatopharyngeus muscles and path-
ostratified columnar ciliated epithelial cells, which is ological changes of the palatinus muscle, associated with
consistent with other studies in horses (Mair et al., chronic denervation (Ducharme and Holcombe, 2001).
1987) and other mammals (Finkelstein et al., 1992). Other structural differences have yet to be investigated.
The oral mucosa consisted of non-keratinised stratified
squamous epithelium, which is the same as other species
(Kuehn and Kahane, 1990; Finkelstein et al., 1992). 6. Conclusion
Lymphoid tissue was found in discrete aggregates in
the rostral soft palate often surrounding mucosal crypts The results of this study have enabled a detailed qual-
at the oral surface. The lymphoid tissue comprised ger- itative and quantitative account of the tissues within the
minal centres and lymphocytes forming the tonsilla veli equine soft palate and will allow for future comparisons
palatini, similar to those described in sheep (Gabriel to be made with horses with palatal dysfunction. The
et al., 2003). Lymphocytes were also found scattered in proportion of different tissue types were found to exhibit
the nasopharyngeal submucosa and respiratory epithe- considerable inter-subject variation. It is possible that
lia. These findings are consistent with previous reports structural differences may predispose some individuals
of the lymphoid tissue within the equine nasopharynx to nasopharyngeal collapse.
(Mair et al., 1987).
Elastin fibres were found within the nasopharyngeal
submucosa. This has previously been reported by Ahern References
(1994) who described a submucosal grid of ‘‘elastic
bands’’ orientated in a rostro-caudal direction over the Ahern, T.J., 1994. The elastic pharynx. Journal of Equine Veterinary
soft palate and caudolateral pharyngeal walls. The elastin Science 14, 462–463.
 L.E. Richardson et al. / The Veterinary Journal 172 (2006) 78–85 85

Beard, W., 1996. Upper respiratory causes of exercise intolerance. Holcombe, S.J., Derksen, F.J., Stick, J.A., Robinson, N.E., 1998.
Veterinary Clinics of North America (Equine Practice) 12, 440–443. Effect of bilateral blockade of the pharyngeal branch of the vagus
Black, J.B., 1977. The structure of the salivary glands of the human nerve on soft palate function in horses. American Journal of
soft palate. Journal of Morphology 153, 107–117. Veterinary Research 59, 504–508.
Cook, W.R., 1981. Some observations on form and function of the Kuehn, D.P., Folkins, J.W., Cutting, C.B., 1982. Relationships
equine upper airway in health and disease. 1: the pharynx. between muscle activity and velar position. Cleft Palate Journal
Proceedings of the American Association of Equine Practitioners 19, 25–35.
27, 355–389. Kuehn, D.P., Folkins, J.W., Linville, R.N., 1988. An electromyo-
Ducharme, N.G., Holcombe, S.J., 2001. The function of the extrinsic graphic study of the musculus uvulae. Cleft Palate Journal 25, 348–
and intrinsic musculature in stabilizing the upper airways. In: 355.
Proceedings of the 2001World Equine Airways Symposium, CD Kuehn, D.P., Kahane, J.C., 1990. Histologic study of the normal
ROM. human adult soft palate. Cleft Palate Journal 27, 26–35.
Ettema, S.L., Kuehn, D.P., 1994. A quantitative histologic study of the Mair, T.S., Batten, E.H., Stokes, C.R., Bourne, F.J., 1987. The
normal human adult soft palate. Journal of Speech and Hearing histological features of the immune system of the equine respira-
Research. 37, 303–313. tory tract. Journal of Comparative Pathology 97, 575–585.
Finkelstein, Y., Meshorer, A., Talmi, Y.P., Zohar, Y., Brenner, Y., Martin, B.B., Reef, V.B., Parente, E.J., Sage, A.B., 2000. Causes of
Gal, R., 1992. The riddle of the uvula. Otolaryngology – Head and poor performance of horses during training, racing or showing: 348
Neck Surgery 107, 444–450. cases (1992–1996). Journal of the American Veterinary Medicine
Franklin, S.H., Naylor, J.R., Lane, J.G., 2002. Effect of dorsal Association 216, 554–558.
displacement of the soft palate on ventilation and airflow during Metternich, F.U., Brusis, T., Koebke, J., Wenzel, S., 2000. Studies of
high intensity exercise. Equine Veterinary Journal Supplement 34, the histomorphology and function of the uvula. Laryngorhinotol-
379–383. ogie 79, 465–470.
Friberg, D., Ansved, T., Borg, K., Carlsson-Nordlander, B., Larsson, Moon, J.B., Thompson, S.A., Jaeckel, E., Canady, J.W., 1998. Muscle
H., Svanborg, E., 1998. Histological indications of a progressive fibre type distribution in the normal human levator veli palatini
snorers disease in upper airway muscle. American Journal of muscle. Cleft Palate-Craniofacial Journal 35, 419–424.
Critical Care Medicine 157, 586–593. Morris, E.A., Seeherman, H.J., 1990. Evaluation of upper respiratory
Gabriel, A., Cocquyt, G., Van Den Broeck, W., 2003. Aspects tract function during strenuous exercise in racehorses. Journal of
anatomique et histologique des tonsilles du mouton. Annales de the American Veterinary Medicine Association 196, 431–438.
Médecine Vétérinaire 147, 251–258. Negus, V.E., 1949. In: Hafner, W. (Ed.), The Comparative Anatomy
Hall, H.D., Schneyer, C.A., 1977. Functional mediation of compen- and Physiology of the Larynx. Heinmann Ltd., London.
satory enlargement of the parotid gland. Cell and Tissue Research Sisson, S., 1975. Equine digestive system. In: Getty (Ed.), Sisson and
184, 249–254. GrossmanÕs Anatomy of Domestic Animals, fifth ed. WB Saunders
Heffron, C.J., Baker, G.J., 1979. Observations on the mechanism of Co., Philadelphia, pp. 471–475.
functional obstruction of the nasopharyngeal airway in the horse. Wells, H., Peronace, A.A.V., 1967. Functional hypertrophy and
Equine Veterinary Journal 11, 142–147. atrophy of the salivary glands of rats. American Journal of
Holcombe, S.J., Derksen, F.J., Stick, J.A., Robinson, N.E., 1997. Physiology 212, 247–251.
Effect of bilateral tenectomy of the tensor veli palatini muscle on Woodson, B.T., Garancis, J.C., Toohil, R.J., 1991. Histopathologic
soft palate function in horses. Journal of the American Veterinary changes in snoring and obstructive sleep apnoea syndrome.
Medicine Association 58, 317–321. Laryngoscope 101, 1318–1322.