From: Subject: Prion Infection of Skeletal Muscle Cells and Papillae in the Tongue -- Mulcahy et al. 78 (13): 6792 -- The Journal of Virology Date: Tue, 4 Apr 2006 21:23:20 -0600 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_0000_01C6582D.FF00E860" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.2180 This is a multi-part message in MIME format. ------=_NextPart_000_0000_01C6582D.FF00E860 Content-Type: text/html; charset="Windows-1252" Content-Transfer-Encoding: quoted-printable Content-Location: http://jvi.asm.org/cgi/content/full/78/13/6792 Prion Infection of Skeletal Muscle Cells and Papillae = in the Tongue -- Mulcahy et al. 78 (13): 6792 -- The Journal of = Virology
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Journal of Virology, July 2004, p. 6792-6798, Vol. 78, No.=20 13
0022-538X/04/$08.00+0     DOI:=20 10.1128/JVI.78.13.6792-6798.2004
Copyright =A9 2004, American Society for Microbiology. All = Rights=20 Reserved.

Prion Infection of Skeletal Muscle Cells and Papillae in the Tongue=20

Ellyn R. Mulcahy,1,3D{dagger}=20 = Jason C.=20 Bartz,1 Anthony E. Kincaid,2 and Richard A.=20 Bessen3* =

Department of Medical Microbiology and Immunology,1 = Department of=20 Physical Therapy, Creighton University, Omaha, Nebraska = 68178,2=20 Department of Veterinary Molecular Biology, Montana State University = Bozeman,=20 Montana 597173

Received 29 September 2003/ Accepted 3 March 2004


3D"   =20 ABSTRACT
3D"Top
3D"Abstract
3D"Introduction
3D"Materials and Methods
3D"Results
3D"Discussion
3D"References
 
T= he=20 presence of the prion agent in skeletal muscle is thought to = be due=20 to the infection of nerve fibers located within the muscle. = We report=20 here that the pathological isoform of the prion protein,=20 PrPSc, accumulates within skeletal muscle cells, = in=20 addition to axons, in the tongue of hamsters following intralingual = and intracerebral inoculation of the HY strain of the = transmissible=20 mink encephalopathy agent. Localization of PrPSc to the = neuromuscular junction suggests that this synapse is a site = for prion=20 agent spread between motor axon terminals and muscle cells.=20 Following intracerebral inoculation, the majority of = PrPSc=20 in the tongue was found in the lamina propria, where it was=20 associated with sensory nerve fibers in the core of the = lingual=20 papillae. PrPSc staining was also identified in = the=20 stratified squamous epithelium of the lingual mucosa. These = findings=20 indicate that prion infection of skeletal muscle cells and = the=20 epithelial layer in the tongue can be established following = the=20 spread of the prion agent from nerve terminals and/or axons = that=20 innervate the tongue. Our data suggest that ingestion of meat = products containing prion-infected tongue could result in = human=20 exposure to the prion agent, while sloughing of = prion-infected=20 epithelial cells at the mucosal surface of the tongue could = be a=20 mechanism for prion agent shedding and subsequent prion = transmission=20 in animals.


3D"   =20 INTRODUCTION
3D"Top
3D"Abstract
3D"Introduction
3D"Materials and Methods
3D"Results
3D"Discussion
3D"References
 
I= n prion=20 diseases, the prion agent is primarily found in the = lymphoreticular=20 and nervous systems (8, 23). In = both=20 natural and experimental prion infections, the prion agent is = commonly amplified by replicating in the lymphoreticular = system=20 prior to entry into nerve cells and subsequent invasion of = the=20 brain (1, 18, 23). = Spread within=20 the nervous system can occur by transport along axons and = between=20 synaptically linked neurons. Experimental studies have = demonstrated=20 that the initial pattern of prion agent spread in the spinal = cord and=20 brain follows defined autonomic, sensory, and motor pathways = (3, 4, 12, 13, 27).=

Modest levels of prion agent replication in skeletal muscle = have=20 been reported in a few studies following intracerebral or = extraneural=20 inoculation of the prion agent. Prion infectivity in skeletal = muscle=20 was first demonstrated in mink with transmissible mink = encephalopathy=20 (TME); the amount of infectious agent in skeletal muscle was=20 10,000-fold less than the amount found in brain (26). = Prion=20 infectivity or accumulation of the abnormal form of the prion = protein, PrPSc, in skeletal muscle has more = recently been=20 described in experimental prion infection of rodents and in = sporadic=20 Creutzfeldt-Jakob disease (CJD) of humans (14),= =20 but the cellular location of agent deposition and/or = replication=20 in muscle has not been identified (3, 6, 40). In = natural=20 prion diseases, PrPSc is found in the peripheral = nervous=20 system of scrapie-infected sheep and goats, and it has been = proposed=20 that it is the peripheral nerves which transverse skeletal=20 muscle that are the source of prion infectivity in muscle (16, 17, = 19, 20). = Attempts to=20 measure the prion agent in muscle of goats infected with the = scrapie=20 agent (18, 20) and = cattle=20 infected with the bovine spongiform encephalopathy (BSE) = agent (10, 42) = have not detected prion infectivity. However, a murine = bioassay=20 was used to measure prion infectivity in these studies, and=20 this assay cannot detect below 104.1 lethal median = doses=20 (LD50) of the BSE agent (10, 42). = Therefore,=20 low levels of prion agent that could be present in muscle = tissue of=20 domestic livestock would not be found using this bioassay. = Routine=20 testing for PrPSc in brains from cattle in Europe = and in=20 lymph nodes from deer and elk in the United States has been=20 implemented to reduce human exposure to BSE and chronic = wasting=20 disease by removing infected animals from the food supply. = However,=20 human consumption of skeletal muscle from a prion-infected = animal is=20 currently considered a low risk for contracting a prion = disease.=20

In this study, we report that in hamsters infected with the = TME=20 agent, by either an intralingual or intracerebral route,=20 PrPSc can accumulate in skeletal muscle cells of the=20 tongue. Localization of PrPSc to the neuromuscular = junction (NMJ) indicates that this synapse can be a site for = prion=20 agent replication and may provide a path by which the prion = agent=20 spreads between skeletal muscle cells and axon terminals.=20 PrPSc was also found in the lingual papillae of = the=20 tongue, specifically in nerve fibers of the lamina propria = and in the=20 stratified squamous epithelium of the tongue surface. These = findings=20 have public health implications and suggest that (i) = ingestion of=20 prion-infected tongue could be a source of prion exposure for = humans,=20 since livestock tongue is not included in the specified risk=20 material; and (ii) normal shedding of the outer layer of the = tongue=20 epithelium can release the prion agent into saliva, which may = be a=20 source for prion agent transmission in livestock and = cervids.=20


3D"   =20 MATERIALS AND METHODS
3D"Top
3D"Abstract
3D"Introduction
3D"Materials and Methods
3D"Results
3D"Discussion
3D"References
 
<= STRONG>Animal=20 inoculations. All procedures involving animals were approved by = the=20 Creighton University Institutional Animal Care and Use = Committee and=20 were in compliance with the Guide for the Care and Use of=20 Laboratory Animals (31). = Hamsters were=20 inoculated in the lingual muscles of the tongue or the = cerebrum with=20 20 =B5l of a brain homogenate from a TME-infected hamster = which=20 contained 107.5 intracerebral LD50 per = ml of=20 the HY TME agent. Following inoculation with the HY TME = agent,=20 hamsters were observed three times per week for the onset of = clinical=20 symptoms. Hamsters were euthanized at either specific = intervals=20 postinfection or after the onset of disease. In the latter = case,=20 animals were culled at 75 to 85 days postinfection for the=20 intracerebral inoculation group and at 95 to 105 days for the = intratongue inoculation group. Tongue and brain were = collected for=20 PrPSc analysis by Western blotting and=20 immunohistochemistry. At least two independent mock-infected = and HY=20 TME-infected tissue samples were used for each of these = analyses, but=20 three or more tissue samples were typically used in each = study.=20 Western blotting and immunohistochemistry results were = similar among=20 individual animals within each control and experimental = group.=20

Tissue preparation for PrPSc Western = blotting.=20 PrPSc was enriched from tongue prior to Western blotting = by=20 tissue extraction in detergent and differential = ultracentrifugation=20 as previously described (3, 4).=20 PrPSc-enriched samples were digested with = proteinase K and=20 resuspended in sodium dodecyl sulfate-polyacrylamide gel=20 electrophoresis (SDS-PAGE) sample loading buffer. = Five-percent brain=20 homogenates were also digested with proteinase K and prepared = for=20 Western blotting as previously described (4). = SDS-PAGE and=20 Western blot analysis were performed using monoclonal = antibody 3F4=20 hybridoma supernatant (22) = (a=20 gift of Victoria Lawson, NIH Rocky Mountain Labs, Hamilton,=20 Mont.) as previously described. Quantification of PrPSc = signal in Western blot assays was performed using ImageQuant=20 software as previously described (4). =

PrPSc immunohistochemistry, immunofluorescence, = and=20 confocal microscopy. Immunostaining for PrPSc in = tongue was=20 performed as previously described (3). = Briefly,=20 animals were intracardially perfused with=20 paraformaldehye-lysine-periodate fixative followed by postfixation = in=20 paraformaldehye-lysine-periodate for 5 h. Paraffin-embedded = tissue=20 sections (4 =B5m) were subjected to antigen retrieval by = pretreatment=20 with formic acid for 20 min. A minimum of 10 tissue sections = per=20 animal was examined for each antibody staining procedure.=20 PrPSc was detected by incubation with monoclonal = 3F4=20 hybridoma supernatant (1:600 dilution) and the ABC-horseradish=20 peroxidase Elite staining method (Vector Laboratories, = Burlingame,=20 Calif.). For PrPSc staining in the double=20 immunofluorescence procedure, the ABC-horseradish peroxidase=20 incubation step in the ABC method was replaced by incubation = with an=20 Alexa Fluor 488 streptavidin conjugate (Molecular Probes, = Portland,=20 Oreg.) at a 1:400 dilution. PrPSc = immunofluorescence=20 staining was combined with rabbit polyclonal antibody = staining for=20 either desmin (1:50 dilution; DAKO Corp., Carpinteria, = Calif.),=20 synaptophysin (1:50 dilution; DAKO Corp.), S100 (1:800 = dilution; DAKO=20 Corp.), cytokeratin (1:800 dilution; DAKO Corp.), or = calcitonin=20 growth-related peptide (CGRP; 1:125 dilution; Peninsula Labs, = Inc.,=20 San Carlos, Calif.). This latter group of primary antibodies = was=20 visualized by incubation with anti-rabbit antibody conjugated = to=20 Alexa Fluor 568 (Molecular Probes). The nuclear counterstain = TO-PRO3=20 (Molecular Probes) was applied to some tissue sections at a=20 concentration of 0.25 =B5M for 10 min. Tissue sections were = mounted=20 with Vectashield Antifade (Vector Laboratories). Images were=20 visualized with epifluorescence using a Bio-Rad Radiance 2000 = confocal system attached to a Nikon Eclipse 800 = microscope.=20


3D"   =20 RESULTS
3D"Top
3D"Abstract
3D"Introduction
3D"Materials and Methods
3D"Results
3D"Discussion
3D"References
 
I= noculation=20 of the HY TME agent into the lingual muscles of the tongue = and the=20 cerebrum resulted in incubation periods of 83 =B1 3 days and = 62 =B1 2=20 days, respectively, in Golden Syrian hamsters (mean =B1 = standard error=20 of the mean). At the onset of clinical symptoms, both groups = had=20 evidence of PrPSc deposition in the tongue (Fig. = 1). = Quantification=20 of the PrPSc polypeptide bands using ImageQuant = software=20 indicated that the amount of PrPSc in the tongue = was=20 approximately fourfold higher following intratongue = inoculation than=20 that following intracerebral inoculation. However, the amount = of=20 PrPSc in the brain was approximately 1,000-fold = higher=20 than that found in comparable amounts of TME-infected tongue=20 following either route of inoculation (Fig. 1).=20 Immunohistochemistry revealed that PrPSc was = located in=20 three regions of the tongue, including the skeletal muscle = (Fig. 2B), the = lamina=20 propria of the papillae (Fig. 2C), and = the=20 stratified squamous epithelium of the tongue mucosa (see = below).=20



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  		FIG. 1. PrPSc = deposition in=20 tongue and brain following inoculation of the HY TME agent. = Hamsters=20 were inoculated with the TME agent or a normal brain = homogenate=20 (mock) by either the intratongue or intracerebral route. = Tongue and=20 brain tissues were collected from animals that were = sacrificed after=20 the onset of clinical disease as described in the text.=20 PrPSc in brain homogenates or = PrPSc-enriched=20 preparations from tongue digested with proteinase K were = prepared as=20 described in Materials and Methods and analyzed by SDS-PAGE = and PrP=20 Western blotting. The amount of tissue analyzed in each lane = is=20 indicated in milligram tissue equivalents (mg eq), and the = molecular=20 masses of polypeptides in kilodaltons are indicated to the = left of=20 the panel. =

 = ;


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  		FIG. 2. PrPSc = deposition in=20 tongue following inoculation of the HY TME agent. Hamsters = were=20 inoculated with a normal brain homogenate (A) or the TME = agent (B=20 and C) by either the intratongue (A and B) or intracerebral = (C)=20 route. Tongues were collected from animals that were = sacrificed=20 after the onset of clinical disease and prepared for=20 immunohistochemistry as described in Materials and Methods.=20 PrPSc immunohistochemistry illustrated = PrPSc=20 (red punctate staining) in skeletal muscle cells (B) and the = lamina=20 propria (C) but an absence of immunostaining in = mock-infected tongue=20 (A). The tissue was counterstained with hematoxylin. The = lamina=20 propria (LP) and stratified squamous epithelium (SSE) of the = tongue=20 are indicated. Bar, 10 =B5m. =

 = ;
Confocal=20 microscopy was used to localize PrPSc to specific cell=20 types in the tongue. Following intratongue inoculation of the = HY TME agent, the majority of PrPSc staining was found=20 within skeletal muscle cells. This was demonstrated in the=20 tongues of clinically ill hamsters at greater than 95 days=20 postinfection by colocalization of desmin staining (i.e., an=20 intermediate filament protein in skeletal muscle cells) in=20 PrPSc-positive cells (Fig. 3A to C). = While=20 PrPSc was widely distributed within skeletal = muscle cells,=20 it was often present in higher amounts at the periphery of = the cell=20 (Fig. 3A). At 1 = week postinoculation of the tongue, PrPSc staining = was=20 found in less than three skeletal muscle cells per tissue = section;=20 the number of PrPSc-positive cells as well as the = amount=20 of PrPSc staining progressively increased in = skeletal=20 muscle cells during the course of subclinical TME infection = (data not=20 shown). At 95 days postinoculation, when hamsters were = clinically=20 affected, skeletal muscle cells throughout the tongue were=20 PrPSc positive. It is worth noting that=20 PrPSc-positive muscle cells were located between=20 PrPSc-negative cells (Fig. 2B and 3A). = These results=20 demonstrate that the TME agent can accumulate within skeletal = muscle=20 cells and suggest that replication can also occur in these = cells. At=20 the clinical stages of TME, a lower amount of = PrPSc=20 staining was found in skeletal muscle cells of the tongue in = hamsters=20 that were intracerebrally inoculated compared to those = inoculated=20 into the tongue with the HY TME agent (data not shown). This = finding=20 was consistent with PrPSc Western blotting data of = tongue=20 from intracerebral- and intratongue-inoculated hamsters (Fig. = 1). In=20 mock-infected controls, specific binding of anti-PrP = monoclonal 3F4=20 antibody was not observed (Fig. 2A). =



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  		FIG. 3. Confocal microscopy of=20 PrPSc in skeletal muscle cells and axons. = Hamsters were=20 inoculated in the lingual muscles with the HY TME agent. At = the=20 onset of clinical symptoms, the tongue was prepared for=20 immunofluorescence staining and confocal microscopy as = described in=20 Materials and Methods. Following immunofluorescence staining = for=20 identification of PrPSc with Alexa Fluor 488 (A, = D, and=20 G), tissue sections were also stained for either desmin (B), = S100=20 (E), or synaptophysin (H) with goat anti-rabbit = immunoglobulin=20 conjugated to Alexa Fluor 568. To investigate the = colocalization of=20 PrPSc with desmin (C), S100 (F), and = synaptophysin (I),=20 green and red fluorescent images were merged. Bar, 10 =B5m. = The insets=20 at the lower left corner (C, F, and I) are two- to threefold = enlargements of smaller boxes in each of the corresponding = panels. =

 = ;
Previously,=20 we demonstrated that PrPSc was associated with axons = in=20 nerve fascicles in the tongue (3). In = the present=20 report, PrPSc staining in nerve fascicles was = demonstrated=20 after 95 days postinfection following intratongue inoculation = of HY=20 TME, but it did not colocalize with S100 staining, which was=20 used as a marker for Schwann cells (Fig. 3D to F). = This=20 finding suggested that the majority of PrPSc was = located=20 within the axons in this experimental model and was less = likely to be=20 within the Schwann cells that surround axons in the = peripheral=20 nervous system. Both large nerve fascicles that travel = between=20 skeletal muscle fascicles and small nerve fascicles located = in the=20 lamina propria were found to contain punctate = PrPSc=20 staining following intratongue inoculation.

The spatial relationship between PrPSc and the NMJ was=20 investigated following intratongue inoculation of the HY TME = agent,=20 since PrPSc was found in both skeletal muscle = cells and=20 axons in the tongue. An antibody to synpatophysin, a marker = for=20 synaptic vesicles in the axon terminal, was used for = identification=20 of NMJs in hamster tongue from clinically ill hamsters at 95=20 days postinfection. Sixty-four percent of = synaptophysin-positive=20 skeletal muscle cells were also PrPSc positive;=20 colocalization of synaptophysin and PrPSc was = found in 25%=20 of these cells (Fig. 3G to I). = In the=20 example illustrated (Fig. 3G to I), = PrPSc staining was localized to the NMJ, = indicating that=20 this is one potential site for PrPSc accumulation = in=20 skeletal muscle cells and/or axon terminals.

The majority of PrPSc staining in the lingual mucosa was=20 located in the lamina propria following intracerebral = inoculation=20 of HY TME, especially within the connective tissue core of = the=20 lingual papillae (Fig. 2C). To = investigate=20 the spatial relationship between PrPSc and nerve = fibers in=20 the lamina propria, we examined PrPSc and CGRP = staining in=20 the tongue at 75 to 85 days postinfection following = intracerebral=20 inoculation of the HY TME agent. CGRP staining in the lamina = propria=20 was used to identify a subset of sensory nerves, since not = all=20 sensory fibers are CGRP positive. PrPSc = immunostaining was=20 identified in 35% of CGRP-positive papillae, but = colocalization of=20 CGRP and PrPSc was found in only 5% of = CGRP-positive=20 papillae (Fig. 4A to=20 C). In 4% of the lingual papillae, PrPSc = staining was=20 also found in the stratified squamous epithelium of the = tongue mucosa=20 following intracerebral inoculation (Fig. 4D). In = these=20 examples, the PrPSc and cytokeratin staining were = in close=20 proximity in the papillary epithelium (Fig. 4D to F); = cytokeratin immunohistochemistry was used as a marker for the = stratified squamous epithelial cells of the tongue. However, = there=20 was no direct overlap of the two staining patterns (Fig. 4F, = inset). A=20 greater amount of PrPSc staining was found = associated with=20 tongue papillae following intracerebral inoculation compared = to=20 inoculation of the HY TME agent into the lingual muscles. No=20 PrPSc-specific staining was observed in the tongue = of=20 mock-infected hamsters (Fig. 4G to = I).=20



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  		FIG. 4. Confocal microscopy of=20 PrPSc in sensory nerve fibers and the stratified = squamous=20 epithelium. Hamsters were inoculated in the cerebrum with = the HY TME=20 agent (A to F) or a normal brain homogenate (G to I) (mock=20 infected). At the onset of clinical symptoms, the tongue was = prepared for confocal microscopy as described in Materials = and=20 Methods. Following immunofluorescence staining for = PrPSc=20 with Alexa Fluor 488 (A, D, and G), tissue sections were = stained for=20 either CGRP (B) or cytokeratin (E and H) with goat = anti-rabbit=20 immunoglobulin conjugated to Alexa Fluor 568. The stain = TO-PRO3 was=20 used to visualize nuclei under blue fluorescence. To = investigate the=20 colocalization of PrPSc with CGRP (C) and = cytokeratin (F=20 and I), green and red fluorescent images were merged. Bar, = 10 =B5m.=20 The insets at the bottom (C and F) are two- to threefold=20 enlargements of smaller boxes in each of the corresponding = panels. =

 = ;
PrPSc=20 staining in the tongue mucosa did not appear in cell groups = that had=20 the morphology of the lingual tonsils. However, we cannot = exclude the=20 possibility that PrPSc was associated with diffuse = lymphoid tissue that is found in the tongue of some = species.=20


3D"   =20 DISCUSSION
3D"Top
3D"Abstract
3D"Introduction
3D"Materials and Methods
3D"Results
3D"Discussion
3D"References
 
I= n this=20 study we report that the prion agent is located within = skeletal=20 muscle cells following inoculation of the HY TME agent into = either=20 the tongue or the cerebrum. While the TME agent was = distributed=20 throughout the individual muscle cells, PrPSc was = also=20 localized to the NMJ in 25% of synaptophysin-positive muscle = cells.=20 These findings do not distinguish whether the = PrPSc=20 staining was present in the presynaptic axon terminal, = postsynaptic=20 skeletal muscle, or in both locations in the NMJ. = PrPSc=20 was also identified in sensory axons of the lamina propria, = mainly in=20 the core of the lingual papillae, but also within the = stratified=20 squamous epithelium. This tissue distribution of the prion = agent in=20 skeletal muscle and oral mucosa has not been previously=20 reported.

Prior studies have demonstrated PrPSc accumulation in the=20 tongue (3, 40) or = prion=20 infectivity in skeletal muscle (6, 26, 40),= =20 but PrPSc staining by immunomorphological methods has = only=20 been reported in nerve fibers that transverse skeletal muscle = (3). = In=20 the present study, PrPSc deposition was found in nerve = fibers=20 and skeletal muscle cells of the tongue. The normal isoform = of=20 the prion protein, PrPC, has been demonstrated to spread = within nerve fibers by axonal transport (5) and is = located=20 in synapses, including subsynaptic areas in both the = presynaptic=20 and postsynaptic cells of the NMJ (2, 15). = Since=20 PrPC is required for PrPSc formation = (7), we = propose that=20 the NMJ is a site for PrPSc formation and/or=20 PrPSc spread between skeletal muscle cells and = axon=20 terminals. In this case, PrPSc spread to, and = accumulation=20 in, skeletal muscle cells in the tongue following = intracerebral=20 inoculation could be due to TME agent replication in the = brain and=20 subsequent anterograde transport within the hypoglossal = nerve, which=20 provides motor innervation to the lingual muscles. = PrPSc=20 may spread across the NMJ and enter skeletal muscle cells, = where=20 additional TME agent replication can occur, since = PrPC is=20 also expressed in skeletal muscle cells (15, 29).= =20 This hypothesis is supported by studies that describe (i) the = spread of the prion agent in both the anterograde and = retrograde=20 directions between synaptically linked nerve cell groups in = the=20 central nervous system (4, 12, 13, 27), = (ii) the=20 retrograde transport of the HY TME agent from the tongue to = the brain=20 in the hypoglossal nerve following intratongue inoculation = (3), = and=20 (iii) prion infectivity in skeletal muscle of transgenic mice = that=20 express PrPC in myocytes (6). = Although the=20 NMJ is morphologically distinct from synapses in the central=20 nervous system, our findings are consistent with this = peripheral=20 synapse serving as a pathway for intercellular spread of the = prion=20 agent between motor axon terminals and muscle cells. =

PrPSc deposition in the lamina propria below the mucosal=20 epithelium was associated with sensory nerve fibers; however, = we=20 cannot exclude the possibility that other types of cells or = axons=20 are infected with the TME agent, since only a portion of=20 PrPSc staining colocalized with CGRP staining. = Given that=20 CGRP staining only labels a subset of sensory fibers, it is = possible=20 that other types of sensory fibers located in the lamina = propria also=20 contain PrPSc. Based on the location of = PrPSc=20 staining and morphology of the adjacent tissue, there was no = evidence=20 that PrPSc was present in the lingual tonsils, = which are=20 also a potential target for TME agent replication. However, = from=20 these studies we cannot exclude that the lingual tonsils are = a site=20 of TME agent replication. PrPSc staining in the = lamina=20 propria was often found in the connective tissue in the core = of=20 papillae; this area has a high concentration of sensory = fibers that=20 innervate mechanoreceptors and taste buds in the tongue (30, 35).=20 PrPSc staining in the lamina propria and tongue = epithelium=20 was more pronounced following intracerebral inoculation = compared to=20 intralingual inoculation. This finding could be due to TME = agent=20 replication in the brain and subsequent transganglionic = transport=20 within the tongue-associated sensory cranial nerves. = PrPSc=20 that was localized to cytokeratin-positive cells could also = be=20 explained by TME infection of sensory nerve fibers that = project into=20 the epithelial cell layers of the tongue (11, 25, 32, 38).=20 Alternatively, prion agent could spread to basal = keratinocytes from=20 sensory nerve endings and lead to PrPSc formation, = since=20 keratinocytes also express PrPC (33, 39). = This=20 proposed pathway for prion agent spread to the mucosa along=20 sensory fibers is analogous to the previous reports of spread = of the CJD agent from the brain to the cornea and olfactory=20 mucosa in sporadic CJD (24, 45; P. = Duffy, J.=20 Wolf, G. Collins, A. G. DeVoe, B. Streeten, and D. Cowen, = Letter, N.=20 Engl. J. Med. 290:692-693, 1974). The low level or = absence of=20 the prion agent in the lymphoreticular system in sporadic CJD = (14, 21) = suggests that the optic and olfactory cranial nerves, = respectively,=20 transport the CJD agent from the brain to these mucosal or = exposed=20 surfaces. Ultrastructural studies will be necessary to = further=20 define the location of PrPSc in the lamina propria and=20 sensory nervous system in the tongue and its spatial = relationship=20 to epithelial cells of the tongue.

In natural prion diseases, prion agent transport from the brain = to=20 the tongue could have implications for human food safety and = animal=20 prion transmission. In sheep with scrapie, cattle with BSE, = and deer=20 with chronic wasting disease, the brain stem regions = containing the=20 tongue-associated cranial nerve nuclei have been reported to = be=20 targets for prion infection. In one or more of these = diseases,=20 PrPSc has been found in the hypoglossal nucleus = (9, 37, 43), the = nucleus=20 of the solitary tract (9, 34, = 37, 43), and = several=20 of the trigeminal nuclei (9, 34, 37, 43, = 44). The = ability=20 of the prion agent to establish infection in brain nuclei = that=20 contribute to, or synapse with, the four cranial nerves that=20 innervate the tongue suggests that this could be a pathway = for=20 centrifugal prion agent spread. The tongue is one of the most = densely=20 innervated extraneural tissues in the human body, with a very = high=20 concentration of both motor and sensory axons (28, 30, 35, 36, 41). = Anterograde=20 prion transport within the hypoglossal nerve would be = expected to=20 result in prion infection of the lingual muscles, while=20 transganglionic transport within the trigeminal, facial, and=20 glossopharyngeal nerves could result in prion spread to the = sensory=20 fibers in the tongue. It is noteworthy that prion infectivity = has=20 been found in the trigeminal ganglia of sheep with scrapie = (19) = and=20 cattle with BSE (42), = indicating=20 that centrifugal spread of these prion agents within the = trigeminal=20 nerve can occur and could result in spread to peripheral = sites=20 including the tongue. Since livestock and cervid tongues are = not=20 banned for human consumption, it is possible that humans = could be=20 exposed to animal prion diseases by ingestion of = prion-infected=20 tongue. Attempts to identify prion infectivity in bovine = tongues=20 have not been successful. However, these studies have used an = infectivity bioassay that cannot detect below = 104.1=20 LD50 of BSE prions, which may not detect low = levels of the=20 prion agent (10, 42).= =20 Our findings also have implications for intraspecies = transmission=20 of animal prion diseases. Establishment of prion infection in = the stratified squamous epithelial cells of the tongue, as = suggested=20 by our findings, could result in sloughing of the prion agent = into the saliva, since this tissue is undergoing continual = cell=20 turnover. In this case, animal behaviors resulting in the = exchange=20 of saliva between hosts may play a role in prion = transmission.


3D"   =20 ACKNOWLEDGMENTS
 
This = work was=20 supported by grants from NIH NIAID (1RO1 AI055043-01), NIH = NCRR=20 (RR15635), and USDA/CREES/NRICGP (2002-35204-12584).

Special thanks are given to Maria Christensen and Emily Hansen = for=20 excellent technical assistance.


3D"   =20 FOOTNOTES
 
* Corresponding author. Mailing address: = Department=20 of Veterinary Molecular Biology, P.O. Box 173610, Montana State = University,=20 Bozeman, MT 59717. Phone: (406) 994-1563. Fax: (406) 994-4303. E-mail: = rbessen{at}montana.edu . 3DBack

3D{dagger}=20 Present = address:=20 Department of Anatomy and Cell Biology, University of Kansas = Medical=20 Center, Kansas City, KS 66160. 3DBack


3D"   =20 REFERENCES
3D"Top
3D"Abstract
3D"Introduction
3D"Materials and Methods
3D"Results
3D"Discussion
3D"References
&nb= sp;

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Journal of Virology, July 2004, p. 6792-6798, Vol. 78, = No.=20 13
0022-538X/04/$08.00+0     DOI:=20 10.1128/JVI.78.13.6792-6798.2004
Copyright =A9 2004, American Society for Microbiology. All = Rights=20 Reserved.


This article has been cited by other=20 articles: (Search Google Scholar for Other Citing Articles)=20

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