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Bull Vet Inst Pulawy 56, 411-414, 2012 MORPHOMETRIC COMPARATIVE STUDY OF THE STRIATUM AND GLOBUS PALLIDUS OF THE COMMON SHREW, BANK VOLE, RABBIT, AND FOX BARBARA WASILEWSKA, JANUSZ NAJDZION, MACIEJ RÓWNIAK, KRYSTYNA BOGUS-NOWAKOWSKA, JACEK J. NOWAKOWSKI,1 AND ANNA ROBAK Department of Comparative Anatomy, 1Department of Ecology and Environmental Protection, University of Warmia and Mazury, 10-727 Olsztyn, Poland [email protected] Received: December 5, 2011 Accepted: August 30, 2012 Abstract The morphology of the striatum (St, caudoputamen complex) and globus pallidus (GP) was studied by stereological methods in representatives of four mammalian orders (Insectivora, Rodentia, Lagomorpha, Carnivora). The aim of our study was to give the first detailed morphometric characteristics of the St and GP in the animals. The paraffin-embedded brain tissue blocks were cut in the coronal plane into 50 µm sections, which were stained for Nissl substance. The morphometric analysis of the St and GP has included such parameters as the volume, numerical density, and total number of neurons. The increase in the volume of the St and GP was accompanied by an increase in the total number of neurons and a decrease in their numerical density. The percentage contribution of the GP volume in the corpus striatum shows progressive traits in the common shrew and fox. Key words: mammals, striatum, globus pallidus, morphometric analysis The striatum (St, or caudoputamen complex, or dorsal striatum) and globus pallidus (GP or GPe in primates) belong to the mammalian basal ganglia. The St and GP are defined as the corpus striatum. A morphometric study of the mammalian St and GP has a long tradition and is related with different quantitative aspects. Data on the volumes of the brain and various brain parts in insectivores and primates has been published by Stephan et al. (16). Age changes in the neuron density, the area occupied by neurons, and the absolute number of cells were investigated by Schröder et al. (14) and Eggers et al. (1) for the whole neuron population. The most extensive volumetric studies on the St and/or GP were mainly carried out in the human (7, 14, 17). The volumetric studies of the St and GP were carried out in schizophrenia (4) and in the attention deficit hyperactivity disorder (18). In the available literature, to our knowledge, there are few comparative studies analysing the morphometric structure of the nuclei of the corpus striatum belonging to different orders of mammals (15, 16). In this study, the St and GP in representatives of four different mammalian orders were morphometrically compared. Knowledge about the number of cells and exact sizes of St and GP could be supported by selective labelling methods such as immunocyto- and immunohistochemistry. It would be also useful to have a validated morphometric model of the St and also GP, which would help in understanding parallels and differences among species. Material and Methods The studies were carried out on the telencephalons of four species of mammals: the common shrew (Sorex araneus – Insectivora, b.w. 11-12 g), bank vole (Clethrionomys glareolus – Rodentia, b.w. 19-24 g), rabbit (Oryctolagus cunniculus – Lagomorpha, b.w. 3.5-4.5 kg), and fox (Vulpes vulpes – Carnivora, b.w. 810 kg). All experimental procedures were approved by the Local Ethics Commission. All animals were given intraperitoneally a lethal dose of 80 mg/kg of Nembutal (Lundbeck, Denmark) and then they were decapitated. Paraffin blocks of the brains were cut into 50 µ m sections in the coronal plane. The sections were stained for Nissl substance. Computer reconstructions of the St and GP. All sections were analysed with a calibrated image analysis system that consisted of a computer equipped with morphometric software (Multi-Scan 8.2, Computer Scanning Systems, Poland) and a light microscope coupled with a digital camera (CM40P, VideoTronic, Germany). Partial microscopic images (512 -512 pixel size) of single brain sections were digitally recorded and then all these images were joined into one large digital 412 section that represented the whole corpus striatum and the adjoining structures. All sections were taken into account in the common shrew, every second section in the bank vole, every fourth section in the rabbit, and eighth section in the fox. Morphometric analysis. On each digital section the boundaries of the individual nuclei were outlined. With the help of IGL Trace software, the 2-D outlines were transformed into 3-D slabs. To evaluate the volume of the St and GP, the volumes of the slabs were totalled according to the formula proposed by Cavaliero (3, 9). The total volume of the corpus striatum presented in this study was the sum of the volumes of the St and GP. The parts of the internal capsule were almost completely excluded from this structure in the rabbit and fox. In the common shrew and bank vole the largest parts of the internal capsule were only excluded. To evaluate the numerical density of cells in each of the analysed nuclei, the optical dissector method was implemented with the guidelines described by West and Gundersen (19). The total number of neurons in each of the nuclei was calculated by multiplying the volume of the given nucleus by the numerical density of cells in it (19). Statistical analysis. The statistical analysis was performed using CSS: Statistica v. 6.0 (Statsoft, USA). Significant differences of the morphometric parameters of the St and GP were tested in the one-factor model of the analysis of variance (ANOVA) and the Tukey’s tests (post hoc analysis). The comparison of two averages was made by using the Student’s t- test. The level of the statistical significance was set at P=0.05. Results Volume. The analysis of the volume of the St and GP showed relative increases in these structures in the examined mammals (in the fox St was 62 times and GP was 78 times larger in comparison to the common shrew) (Table 1). The St and GP volumes of examined mammals were significantly (P<0.05) different. The percentage contributions of the St volume in the whole volume of the corpus striatum was the largest in the bank vole, and the smallest in the fox (Table 1). To assess the percentage contributions of the St and GP volume, the Tukey’s post hoc analysis was used, which enabled to distinguish three groups of animals. The first group was formed by the common shrew, the second by the bank vole and rabbit, and the third by the fox. Numerical density. The average number of neurons in 1 mm3 of St and GP decreased with the growth of the volume of these structures in the analysed mammals (Table 1). Tukey’s post hoc analysis revealed the presence of three various groups in regard to the numerical density of striatal neurons and also pallidal neurons. The first group constituted the common shrew, the second the bank vole, and the third the rabbit and fox. The presence of the negative correlation between the average numerical density and the average volume of the St (r= −0.61, N=22, P<0.05; Fig. 1) and GP (r= − 0.55, N=22, P<0.05) was ascertained. Fig. 1. The correlation between the average numerical density of the St and the average volume of the St. 413 Fig. 2. The correlation between the average total number of neurons of the St and the average volume of the St. Table 1 The morphometric parameters and the percentages of the volume of the St and GP in the corpus striatum of the studied species Volume of St (mm3) Volume of GP (mm3) Density of St (N/mm3) Density of GP (N/mm3) Number of St cells (N) Number of GP cells (N) Percentages of the volume of St Percentages of the volume of GP Percentage of neurons of St Percentage of neurons of GP Common shrew 1.34 ±0.11 0.15 ±0.02 229,595 ±17,658 52,931 ±6,477 308,035±35,059 8,043 ±1,854 89.88 ±1.34 10.12 ±1.34 97.46 ±0.46 2.54 ±0.46 The total number of neurons. The population of cells of the St and GP was the smallest in the bank vole and the largest in the fox (Table 1). On the basis of Tukey’s post hoc analysis, three different groups were ascertained regarding the total numbers of St and GP neurons. The first of them was composed of the foxes, the second of the rabbits, and the third of the common shrew and bank vole. In the analysed animals, the presence of correlation between the average total number of neurons and the average volume was ascertained in the St (r=0.99, N=22, P<0.05; Fig. 2) and GP (r=0.98, N=22, P<0.05). The presence of the negative correlation between the average number of striatal neurons and the average numerical density of the St (r= -0.60, N=22, P<0.05) was found. In this study we took into consideration the value of the total number of neurons of the whole corpus striatum. We used this value in order to estimate the percentage contributions of the total neuronal population of the St and GP (Table 1). Tukey’s post hoc analysis revealed the presence of two groups of animals for the Bank vole 3.01 ±0.27 0.22 ±0.02 72,641 ±6,078 15,642 ±2,462 218,926±26,929 3,369 ±586 93.32 ±0.30 6.68 ±0.30 98.48 ±0.22 1.52 ±0.22 Rabbit 26.10 ±0.62 2.20 ±0.11 40,462 ±4,731 6,947 ±1,082 1,051,746±150,173 14,993 ±2,599 92.23 ±0.39 7.77 ±0.39 98.57 ±0.31 1.43 ±0.31 Fox 83.21 ±0.22 11.72 ±0.55 26,390 ±1,133 4,123 ±339 2,203,879±99,634 48,746 ±6,079 87.66 ±0.50 12.34 ±0.50 97.84 ±0.24 2.16 ±0.24 percentage contribution of cellular population of the St and GP. The first group was formed by the common shrew and fox and the second by the bank vole and rabbit. Discussion The presented results indicate that the volumes of the St and GP showed a relative increase in the examined mammals, meaning that the smallest volume was found in the common shrew and the largest volume in the fox. However, the comparison of the present volumetric data with the data of other authors is often difficult, because most of those observations were carried out using the miscellaneous methods of measure (2, 16). Stephan et al. (16) included the nucleus accumbens in the St. The most information concerns the volume of St and GP in the man (47, 6, 7, 1414, 17176). The volumetric percentages of the GP in the corpus striatum are the largest in the common shrew and the fox (presented results), which suggests that GP plays a more 414 important role in both these species. This increase probably is a consequence of the tremendous relative enlargement of other brain parts, for instance, of the St in both animals (our study) and of the isocortex in the fox (11). According to Stephan (15), the percentage of the St volume in relation to the telencephalic volume or to the whole brain volume shows a clear decrease in this parameter in insectivores and primates. The percentage of the St volume showed the lowest value in the man (15). In the present study, we observed that with the increment of the volume of the St and GP, the numerical density of the cell population of these structures decreases. The presence of the negative correlation between the volume of given structure and the average numerical density of the cell population in it has previously been described by Kowiański et al. (55), Równiak et al. (1313) in the amygdala, and Najdzion et al. (8) in the medial geniculate body. In the mouse, St neuron-packing density ranges from approximately 50,000 to 100,000 neurons/mm3 (1210). In the bank vole, the representative of the rodents, the numerical density is comparable with this value (72,641.60). In the rat, the value of the numerical density is two times larger than in the bank vole (141,500 neurons/mm3) (1010). However, the numerical density of the GP cells of the rat does not show considerable differences (17,300 neurons/mm3) (1010). In the man, the numerical density of striatal and pallidal neurons has been examined by some authors (77, 1717). The value of this parameter in the man is clearly smaller in comparison with the numerical density of cell population of the analysed structure in the common shrew, bank vole, rabbit, and fox (77, 17). The presented results indicate that the total number of neurons generally showed a relative increase in the examined mammals. An increase in the total number of neurons with an increase in the volume has been shown by Kowiański et al. (55) in the claustrum and Równiak et al. (1313) in the amygdala of mammals. According to Fentress et al. (2) the St of an adult mouse contains approximately 690,000 neurons on each side but in the bank vole it has three times less neurons. The differences in the total number of neurons are probably the result of the differences in size and weight of the body mass in both analysed animals. In the man, the total number of neurons in the St is 68.1 million (66), while in the GP/GPe ranges from 540,000 to 465,000 (1717). The current study provides quantitative data on the volume, numerical density, and total number of neurons of the St and GP in representatives of four mammalian orders. We hope that these results have cognitive, as well as practical value and they may help in understanding the pathophysiology and proper functioning of the nervous system. References 1. Eggers R., Knebel G., Haug H.: Morphometric studies of biological changes in synapses of the human caudate nucleus. Z Gerontol 1991, 24, 302-305. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. Fentress J.C., Stanfield B.B., Cowan W.M.: Observation on the development of the striatum in mice and rats. Anat Embryol 1981, 163, 275-298. Gundersen H.J.G., Jensen E.B.: The efficiency of systematic sampling in stereology and its prediction. J Microsc 1987, 147, 229-263. 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