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Increased Brain Size and Glial Cell Number in CD81-Null Mice

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Increased Brain Size and Glial Cell Number in CD81-Null Mice THE JOURNAL OF COMPARATIVE NEUROLOGY 453:22–32 (2002) Increased Brain Size and Glial Cell Number in CD81-Null Mice ELDON E. GEISERT, JR.,1* ROBERT W. WILLIAMS,1 GRACE R. GEISERT,1 LIYING FAN,1 ANDREW M. ASBURY,1 HOLDEN T. MAECKER,2 JUN DENG,2 AND SHOSHANA LEVY2 1Department of Anatomy and Neurobiology, and Neuroscience Institute, University of Tennessee, Health Science Center, Memphis, Tennessee 38163 2Department of Medicine, Division of Oncology, Stanford University Medical Center, Stanford, California 94305 ABSTRACT A key issue in the development of the central nervous system (CNS) is understanding the molecular mechanisms regulating cell number. The present study examines the role of CD81 (previously known as TAPA, the target of the antiproliferative antibody) in the control of brain size and glial cell number. CD81 is a member of the tetraspanin family of proteins. This group of small membrane proteins is associated with the regulation of cell migration and mitotic activity. Glial cells express CD81, and antibodies directed against this protein suppress the mitotic activity of cultured cells. In this study, we examine the effects of the CD81 Ϫ/Ϫ mutation on the CNS of mature mice. These mice have extremely large brains, as much as 30% larger than the brains of wild-type (ϩ/ϩ) littermates. The increase in brain weight is accompanied by an increase in the number astrocytes and microglia, whereas the number of neurons and oligodendrocytes in the CD81 Ϫ/Ϫ animals appears to be normal. When the CD81 Ϫ/Ϫ mutation is placed on different genetic backgrounds, there is a remark- able range in the penetrance of the null allele phenotype, demonstrating that the mutation can be affected by modifier loci. This work provides support for the role of CD81 in the regulation of astrocyte and microglial number, perhaps by regulating cell proliferation by a contact inhibition-dependent mechanism. J. Comp. Neurol. 453:22–32, 2002. © 2002 Wiley-Liss, Inc. Indexing terms: astroglia; microglia; oligodendrocytes; cell number; tetraspanin The cellular composition of the brain and spinal cord (Ikeyama et al., 1993; Yanez-Mo et al., 1998; Hemler, belies a series of molecular events that control the final 1998; Fitter et al., 1999; Stipp et al., 2001; Charrin et al., number of each cell type within the mature central ner- 2001). These tetraspanin complexes can link events of vous system (CNS). The present study focuses on CD81 cellular adhesion to intracellular signaling cascades (Jen- (previously known as the target of the antiproliferative nings et al., 1990; Yatomi et al., 1993; Schick and Levy, antibody TAPA), a small membrane protein that is ex- 1993; Berditchevski et al., 1997). Like other members of pressed by all glial cells in the CNS. As its original name suggests, antibodies directed against this protein sup- press the mitotic activity of cultured cells (Oren et al., Grant sponsor: PHS; Grant number: RO1 EY12369; Grant number: P30 1990; Geisert et al., 1996). CD81 is a member of the EY13080; Grant number: CA24233; Grant number: AI45900. recently defined tetraspanin family of proteins (http:// Dr. Maecker’s current address is Becton-Dickinson Immunocytometry www.ksu.edu/tetraspan/thepage.htm) that is structurally Systems, 2350, Qume Drive, San Jose, CA 95131. related, with four transmembrane domains and two rela- *Correspondence to: Eldon E. Geisert, Jr., Department of Anatomy and tively small extracellular loops (Jennings et al., 1990; Neurobiology, University of Tennessee, Medical Center, 855 Monroe Ave- Kaprielian and Patterson, 1993; Yatomi et al., 1993; Dong nue, Memphis, TN 38163. E-mail: [email protected] Received 11 December 2001; Revised 5 April 2002; Accepted 21 June et al., 1995; Kopczynski et al., 1996; Tachibana et al., 2002 1997; Todd et al., 1998; Le Naour et al., 2000; Miyado et DOI 10.1002/cne.10364 al., 2000; Serru et al., 2000). Many of these tetraspanins Published online the week of September 16, 2002 in Wiley InterScience are found in association with adhesion molecules (www.interscience.wiley.com). © 2002 WILEY-LISS, INC. BIG BRAINS IN CD81 ؊/؊ MICE 23 this family, CD81 is part of a protein complex within the derived from C57BL/6ByJ and BALB/cByJ. CD81 Ϫ/Ϫ plasma membrane (Takahashi et al., 1990; Bradbury et mice on both inbred backgrounds have difficulty breeding al., 1992; Schick and Levy, 1993; Yanez-Mo et al., 1998; and are propagated as CD81 heterozygotes. We also gen- Stipp et al., 2001). The specific functional role of CD81 is erated CD81 on a hybrid F1 background by mating BALB/ dependent on its molecular context and the cell type it is cByJ and C57BL/6ByJ carriers. All mice used in this study associated with. For example, CD81 can be associated were over 70 days of age. We studied male and female with the control of mitotic activity in lymphocytes (Oren et mice of all three genotypes. The majority of CD81 Ϫ/Ϫ al., 1990), astrocytes (Geisert et al., 1996), or retinal pig- animals were females. There are no statistically signifi- ment epithelium (Geisert et al., 2002). cant sex differences in brain weights of normal adult mice In the CNS, CD81 appears to be expressed by all major belonging to any of the three key background strains glial cell classes: ependymal cells, choroid cells, astrocytes, (129X1/SvJ, C57BL/6ByJ, and BALB/cByJ; data available oligodendrocytes (Sullivan and Geisert, 1998), and micro- on-line at www.nervenet.org/main/databases.html, eight glia (Dijkstra et al., 2000). Hints regarding the functional cases minimum per sex per strain). role of this tetraspanin can be gleaned from its dynamic Mice were anesthetized with a mixture of xylazine (13 temporal/developmental regulation. During CNS develop- mg/kg, Rompun) and ketamine (87 mg/kg, Ketalar), which ment, neurons are born first and most glial cells are born was administered by intraperitoneal injection. The ani- in a later phase (Caviness, 1982). In this latter phase of mals were perfused through the heart with a solution of brain development, astrocytes and oligodendrocytes are 0.01 M phosphate buffered saline (PBS, pH 7.5) followed born and mature (Ling and Leblond, 1973; Caviness, 1982; by 4% paraformaldehyde in 0.1 M phosphate buffer (PB, Parnavelas et al., 1983). Most of these cells are born pH 7.5). Each brain was removed from the skull, and the postpartum in the rodent, with peak mitotic activity oc- spinal cord was separated at a level that corresponds to curring during the first 10 days (Ling and Leblond, 1973; the decussation of the pyramids. The brains were then Parnavelas et al., 1983). By the end of the second postna- blotted dry and weighed on an electronic balance with an tal week, astrocytes mature, down-regulate their mitotic accuracy of Ϯ 1 mg. activity, and form relatively stable populations (Parna- Brains were placed in a 30% sucrose solution and were velas et al., 1983). For oligodendrocytes, the developmen- sectioned at 50 ␮m with a freezing microtome. All sections tal pattern differs from that of the astrocyte. In the second were stored in borate buffered saline (pH 8.4) at 4°C. One postnatal week, there is an overproduction of oligodendro- 1-in-5 series of sections was mounted on glass slides and cytes. Then as myelination occurs, oligodendrocytes un- stained by the Nissl method. A second series of sections was dergo an apoptotic cell death if appropriate contacts with stained for the astrocytic marker glial fibrillary acidic pro- axons are not made (Gard and Pfeiffer, 1989; Hardy and tein (GFAP). Selected sections from each animal were also Reynolds, 1991). It is during this period of decreased glial stained for the following: (1) a second astrocyte marker using mitosis when CD81 is up-regulated dramatically. Relative S-100 (Sigma, St. Louis, MO), (2) neurons using NeuN to the adult, the overall level of this protein increases from (Chemicon, Temecula, CA), (3) oligodendrocytes using RIP 15% at postnatal day (P) 7 to 70% at P14 (Sullivan and (Chemicon), and (4) microglia using Mac-1 (PharMingen, Geisert, 1998). This tight correlation of CD81 expression San Diego, CA). Detailed methods for immunostaining ap- with the down-regulation of glial cell proliferation is con- pear in Sullivan and Geisert (1998). The sections were sistent with the hypothesis that CD81 plays a role in placed in 4% bovine serum albumin (BSA, Sigma) in 0.2 M controlling glial cell number (Sullivan and Geisert, 1998). borate buffered saline (BBS, pH 8.4) for 2 hours, then placed Based on its apparent function in vitro and its develop- in rabbit anti-GFAP (Immunon) 20 ␮g/ml in 0.2 M BBS with mental expression pattern, we consider CD81 to be part of 1% BSA. Sections were washed 3ϫ for 10 minutes in BBS a molecular complex regulating glial cell number. To test and transferred to a solution containing the secondary anti- this hypothesis, we examined the effects of a CD81 Ϫ/Ϫ body (a peroxidase-conjugated donkey anti-rabbit immuno- mutation on the structure of the adult brain. If this pro- globulin G (IgG) or anti-mouse IgG (Jackson Immuno Re- tein is part of the molecular process modulating mitotic search Laboratories Inc.) for 2 hours at room temperature. activity, then we would predict an increase in the number Solutions with primary or secondary antibodies also con- of glial cells in the brains of CD81 Ϫ/Ϫ animal and possi- tained either 0.05% dimethyl sulfoxide or 0.05% Triton bly an overall increase in brain size. X-100. The sections were rinsed in 0.2 M BBS, followed by three rinses in 0.1 M PB (pH 7.2), and incubated in a solution containing 1 mg/ml of 3,3Ј-diaminobenzidine tetrahydrochlo- MATERIAL AND METHODS ride (Sigma) in 0.02 M PB (pH 7.4) and 5 ␮l/ml of 3% All animal protocols were approved by the Animal Care hydrogen peroxide for 15–30 minutes at room temperature.
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