Biomorphometric characteristics of different types of sensilla detected on the antenna of Helicoverpa armigera by scanning electron microscopy

A morphometric study on H. armigera antenna showed four styles of sensilla, i.e., styloconica, chaetica, coeloconica, and trichodea, and their numbers were estimated. Sensilla trichodea detect inter and intraspecific communication signals and was the most numerous. They were divided into three types: type I, the longest, with a length of 34.04 ± 3.16 μm and about 2.16 to 2.42 μm in diameter at its base; 2) type II, intermediate, with a length of 22.58 ± 0.77 μm and basal diameter of 1.8–2.52 μm; 3) type III, the shortest sensilla trichodea, with a length of 7.62 ± 0.4 μm and a range in diameter similar to that of type II. The length of the female sensilla trichodea was longer than that of the male. The total number of sensilla trichodea was estimated to be 7520 on the antenna of the female, and 6831 on the male antenna. The lengths of the sensilla trichodea type I and type III were significantly different on male (t = 4.6881, P = 0.0034) and female antenna (t = 18.9852, P = 0.0001). An estimation of the predicted surface area of the most numerous type I on sampled segments between the 12th and 20th segments from a female of H. armigera showed a surface area of 5 × 10³ μm² and a sensillar density of 38 sensilla/10³ μm². The fraction of sensilla-occupied surface area was 0.4 μm².     

Antennal sensilla of male and female Telenomus reynoldsi gordh and coker (Hymenoptera : Scelionidae)

Sensilla on the antennae of both sexes of Telenomus reynoldsi (Hymenoptera : Scelionidae) were studied with the scanning electron microscope to determine their structure and possible function in mating and host recognition. Two types of setiform sensilla occur: curved, grooved sensilla trichodea and erect, stout sensilla chaetica. Horn-like sensilla trichodea curvata are abundant on the male antennae, but are less abundant and only present dorsally on the female. Sensilla basiconica are unique to the ventral surface of the apical 4 segments of the female antennae. Sensilla styloconica are located on A4 and A10 of the female and A7 and A9 of the male. Multiporous grooved sensilla are present on A4, A6, and A11 of the female and A4 and A6-A11 of the male. A possible sex gland is located on the venter of A5 of the male. The functions of sensilla trichodea curvata and sensilla basiconica in courtship and host recognition, respectively, are hypothesized by correlation with behavioral observations.     

食蚜瘿蚊触角的扫描电镜观察

用扫描电子显微镜对食蚜瘿蚊角角进行了观察.结果显示,雌雄触角都为14节,其中雄性约为2000μm,雌性约为1 050μm.电镜下可观察到食蚜瘿蚊触角有6种类型的感受器,即：刺形感受器、毛形感受器、锥形感受器、腔形感受器、柱形感受器和环丝.刺形感受器较长,约67.5μm,基部有膜状的窝.毛形感受器长约61μm,末端弯曲.锥形感受器呈钉状着生在表皮上,长约4.7μm.腔形感受器呈凹陷状,腔的直径约为1.2μm.柱形感受器着生在雄虫鞭节的第二亚节,长约21μm,直径约为1.5μm.环丝,是瘿蚊类昆虫触角中特殊的结构,它通过着生在一系列腔中的的短梗,连结成环状附着在触角各亚节的表面.刺形和锥形感受器在数量上,雌雄之间差别不大;柱形感受器只在雄虫中发现;雄虫触角上的腔形感受器在数量上要比雌虫多.     

Sexual dimorphism of tarsal receptors and sensory equipment of the ovipositor in the European corn borer,Ostrinia nubilalis

Summary Sensilla on legs and ovipositor of the moth Ostrinia nubilalis were investigated by light and scanning electron microscopy. The ovipositor is composed of two papillae densely packed with medium length mechanoreceptor sensilla (MRb: 80–160 m, n=420–460). Long mechanoreceptor sensilla (MRa: 250–300 m, n=20–24) and contact chemoreceptors (CRa: 30–40 m, n=20–28) are evenly distributed at the periphery of these papillae. Legs support contact chemoreceptors (CRa), scattered among the scales. The pretarsus structure of each leg includes a single contact chemoreceptor (125 m) inserted dorsally. The fifth tarsomere bears a ventral area without scales on which contact chemoreceptors are disposed in two parallel rows (CRb: 40–60 m). A sexual dimorphism was found in the number and density of these sensilla (females: mean = 5.3, SD=1.0; males: mean = 3.3, SD=0.7), and in the size of the sensory field. The possible involvement of these sensory structures in oviposition site selection is discussed.     

Antennal sensilla of the click beetle,Melanotus villosus (Geoffroy) (Coleoptera: Elateridae)

The typology, number and placement of antennal sensilla of the click beetle Melanotus villosus (Geoffroy) (Coleoptera: Elateridae) were studied using scanning electron microscopy. On both the males and females the antennae are made up of the scape, pedicel and nine flagellomeres. Two types of basiconic sensilla, three types of trichoid sensilla, one type of styloconic sensilla, one type of chetoid sensilla, dome-shaped sensilla, grooved pegs, and Böhm sensilla all appear on the antennae of the beetles of both sexes, with the exception of trichoid sensilla type II, whose large number (average of 1635 hairs per antenna) was found only in male beetles. Sensilla trichodea type II evidently respond to the sex pheromone produced by the female beetle. Unlike the other two click beetles, studied up till now, Agriotes obscurus and Limonius aeruginosus, the trichoid and basiconic sensilla of M. villosus, whose proven or assumed function is olfactory, are located predominantly on the flagellomeres ventral extensions. It is assumed that the placement of the olfactory sensilla, mainly on the ventral side of M. villosuss antennae, and their more or less even distribution on the flagellomeres, can be seen as morphological adaptation of this species of insect, whose specific behavioural reaction of olfactory searching is flying, both before and after contact with an odour plume.     

Sexual dimorphism of antennal and ovipositor sensilla of Tetrastichus sp. (Hymenoptera: Eulophidae)

Tetrastichus sp. (Hymenoptera: Eulophidae) is a primary parasitoid of the Metisa plana (Lepidoptera: Psychidae), an oil palm bagworm. The sensilla on the surface of the antenna and ovipositor of Tetrastichus sp. were examined using a scanning electron microscope. The antennae of both male and female Tetrastichus sp. are geniculate in shape and hinged at the scape-pedicel joint. The female antenna is about 200 µm longer than the male antenna. However, the male antenna has an additional flagellomere compared to the female antenna. In total, eight different types of antennal sensilla were observed on the antenna of Tetrastichus sp.: trichoid sensilla type 1, 2, 3, 4, placoid sensilla type 1 and 2, basiconic sensilla, and campaniform sensilla. The antenna of the female Tetrastichus sp. lacks placoid sensilla type 2 and campaniform sensilla. The distribution and abundance of the antennal sensilla were compared between the male and female Tetrastichus sp. and discussed. On the ovipositor stylet of Tetrastichus sp., coeloconic sensilla, styloconic sensilla and campaniform sensilla were observed. Trichoid sensilla were observed at the medial part of the distal extremity of the ovipositor.     

The Antennal Sensilla of Oxelytrum erythrurum (Blanchard) and Oxelytrum apicale (Brullé) (Coleoptera: Silphidae)

The typology and placement of antennal sensilla of the carrion beetles Oxelytrum erythrurum (Blanchard) and Oxelytrum apicalis (Brullé) (Coleoptera: Silphidae) were studied using scanning electron microscopy. Two types of sensilla chaetica, two types of sensilla trichodea, four types of sensilla basiconica, one type of sensilla coeloconica, and an unidentified type of sensillum were found in both species. Sensilla chaetica type 1 are found on the antennomeres proximal to antennal club (A1?CA8); chaetica type 2 are found on the club (A9?CA11). Sensilla trichodea are found on A9?CA11; one type (T1) is found on the proximal portion of the club, the other type (T2) on the apical portion. Basiconica type 1 are found on the dorsal surface of A9?CA11; they are much denser on the apical portion of the antennal club than on the proximal. In O. erythrurum, a nocturnal species of the Chaco-Pampean plain, T2 two are found on A10 and A11. In Oxelytrum apicale, a mountain species, probably diurnal, only A11 bears T2, but they are denser than in the other species. It is suggested that O. apicale depends more on contact chemoreception than O. erythrurum. The ventral surface of the antennal clubs shows no remarkable difference between species.     

Comparative study of the antennal sensilla of five species of root maggots: Delia radicum L., D. floralis F., D. antiqua Mg., D. platura MG. (Diptera : Anthomyiidae) and Psila rosae F. (Diptera : Psilidae)

A comparative study of the antennal sensilla of Delia radicum L., D. floralis F., D. antiqua Mg., D. platura Mg. (Diptera : Anthomyiidae) and Psila rosae F. Diptera Psilidae) is undertaken. For both sexes of each species, the type, distribution, and density of sensilla are determined. All 5 species have trichoid (olfactory) and grooved (olfactory) sensilla. Basiconica I (blunt) sensilla (olfactory) are found on each of the species examined, except D. platura. Basiconica II (tapered) (olfactory) and clavate (olfactory) sensilla are found only on Delia species. Also, only Delia species have single-chambered, dorsal pits, and these contain basiconic II pit sensilla (olfactory). Common to all 5 species is a multi-chambered ventral pit (olfactory). In the ventral pit, all 5 species have grooved pit sensilla (olfactory). In addition to this type of sensillum the Delia species have smooth-walled conical pit sensilla (hygro-/thermosensitive) and P. rosae has granular pit sensilla (hygro-/thermosensitive). Smooth-walled tapered pit sensilla (hygro-/thermosensitive) are found in D. radicum. Similarities and differences in the density of surface sensilla between dorsal and ventral funicular surfaces, male and female flies, and oligophagous (D. antiqua, D. radicum, D. floralis and P. rosae) and polyphagous (D. platura) species are compared. Several differences in sensillum density between the dorsal and ventral funicular surfaces are observed, but these do not fit into a consistent trend. Except for D. radicum, there are differences in sensillum density between male and female flies. For the oligophagous species, females have a greater sensillum density, whilst for the polyphagous D. platura males have a greater sensillum density. Comparisons between species show the greatest differences between the Delia species and P. rosae, and within the 4 Delia species, differences in sensillum density do not correlate with host range or body size.     

Sense organs on the ovipositor of Macrocentrus cingulum Brischke (Hymenoptera: Braconidae): their probable role in stinging,oviposition and host selection process

Parasitoid wasps from the insect order Hymenoptera can be deployed successfully as biological control agents for a number of pests, and have previously been introduced for the control of corn pest insect species from the Lepidopteran genus Ostrinia. Organs on the ovipositor of parasitoid wasps have mechanical and tactile senses that coordinate the complex movements of egg laying, and the ovipositor of Hymenopteran insects have evolved associated venom glands as part of their stinging defense. The ovipositor of parasitic wasps has evolved an additional function as a piercing organ that is required for the deposition of eggs within suitable host larvae. The morphology and ultrastructure of sense organs on the ovipositor and sheath of Macrocentrus cingulum Brischke (Hymenoptera: Braconidae) are described using scanning and transmission electron microscopy. Three types of sensilla trichodea were shown to be abundant on the outer sheath of the ovipositor, with types II and III being most distal, and the inner surface of the ovipositor covered with microtrichia, more densely near the apex. Sensilla coeloconica are distributed on both ventral and dorsal valves, while campaniform sensilla and secretory pores are only located on the dorsal valve. The olistheter-like interlocking mechanism, as well as the morphology of the ventral and dorsal valve tips and the ventral valve seal may be important in stinging, oviposition and in the host selection process.     

斑鞘豆叶甲触角感器的超微结构

斑鞘豆叶甲是大豆苗期重要害虫,本文利用扫描电镜技术观察了斑鞘豆叶甲触角感器超微形态与分布。结果表明:斑鞘豆叶甲触角感器绝大部分着生于鞭节,在鞭亚节端部和末节凹陷区感器分布密集,类型较多。基于感器外部形态可分为8种类型:毛形感器Ⅰ型和Ⅱ型、刺形感器、锥形感器、指形感器、腔锥形感器、栓锥形感器Ⅰ型和Ⅱ型、钟形感器和B hm氏鬃毛。其中毛形感器数量最多,其次是锥形感器,钟形感器最少,仅分布于雄虫触角,还着生有大量表皮孔。雌、雄虫触角感器在形态、数量和分布上均存在差异,雄虫毛形和刺形感器显著长于雌虫,刺形感器端部膨大,两种感器的数量也明显多于雌虫;雌虫与雄虫相比末节背面也具感器密集的凹陷区,指形感器短于雄虫但数量显著多于雄虫。斑鞘豆叶甲触角感器种类丰富,分布密集,雌、雄虫感器存在明显的性二型现象,其结构和类型表现出种间分化特性。     

Fine structure of Chloromyxum menticirrhi n. sp. (Myxozoa) infecting the urinary bladder of the marine teleost Menticirrhus americanus (Sciaenidae) in Southern Brazil

A myxosporidian was found in the urinary bladder of the teleost Menticirrhus americanus Linnaeus, 1758 (Sciaenidae) collected from the South Atlantic coast of Brazil. Polysporic amoeboid plasmodia containing sporoblasts, developing pansporoblasts and spores were free in the bladder lumen. The prevalence of infection was 17.64% (15/85). Unfixed spores were spherical to subspherical, on average 10.5 μm long, 9.8 μm wide and 10.1 μm thick (n=25), and fixed spores measured 10.1×9.5×9.7 μm. The two spore valves were of equal size and each possessed prominent sutural lines and about 41 (37–45) surface ridges aligned parallel with the suture line. These ridges gave transverse sections a cog-wheel-like outline. The spores contained four pyriform polar capsules of equal size (3.20×2.0 μm) (n=25) (fixed), each with a polar filament having 3–4 (rarely 5) coils. The binucleate sporoplasm was irregular in shape, with granular matrix and randomly distributed dense bodies. The shape and dimensions of the spore, as well as the number, position and arrangement of the surface ridges, polar capsules and polar filament indicate that this is a new species, herein designated Chloromyxum menticirrhi. The gill, liver, gall bladder and intestine of the host showed no abnormalities.     

Fine Structure of the Plasmodia and Myxospore of Ellipsomyxa gobioides n. sp. (Myxozoa) Found in the Gallbladder of Gobioides broussonnetii (Teleostei: Gobiidae) from the Lower Amazon River

A fish infecting myxosporean Ellipsomyxa gobioides n. sp. is described in the gallbladder of the Amazonian dragon fish Gobioides broussonnetii. Irregular disporous plasmodia (up to ~30 μm in diameter) with long branched and anastomosed pseudopodia were found attached to the gallbladder wall. Mature ellipsoid myxospores occurring floating in the bile measured 6.8 (6.5–7.0) μm (n = 30) long, 7.2 (6.9–7.5) μm (n = 15) wide, and 13.1 (12.8–13.5) (n = 25) thick. They had smooth thin valves elongated in the direction perpendicular to the plane of the straight central transverse sutural line. The two ellipsoidal polar capsules (PC) opened some distance from the sutural line on opposite sides, each measuring 4.6 (4.3–4.8) μm (n = 15) long and 2.5 (2.1–2.7) μm (n = 20) wide. Distance between PC 3.5 (3.1–3.8) μm (n = 15) in apical view. The polar filament was isofilar and consisted of a single coil with five or six turns. The objective of this study was to characterize this new species based on its morphological differences from the three previously described species. This is the first reported species of genus Ellipsomyxa from among the South American fauna.     

Sensilla of Carabus (Isiocarabus) fiduciarius saishutoicus Csiki (Coleoptera : Carabidae)

Sensilla on the antenna of the ground beetle, Carabus (Isiocarabus) fiduciarius saishutoicus Csiki (Coleoptera: Carabidae), were investigated using scanning electron microscopy. In both sexes, 5 types of sensilla trichodea, 7 types of sensilla basiconica, 1 type of sensilla ampullacea, and 2 types of sensilla campaniformia are distinguished. No sexual differences in the types and distribution of sensilla are found on the antenna, except a ventral, depressed, hairless region of the male. Various types of sensilla present on the mouthparts, head, pronotum, elytron, hindwing, legs, abdominal tergites and sternites, aedeagus, and ovipositor are also described.     

Ultrastructure structure of antennal sensilla of carabid beetle Carabus elysii Thomson, 1856 (Coleoptera: Carabidae)

The sensilla type, number and distribution in male and female adults of Carabus elysii Thomson, 1856 (Coleoptera: Carabidae) were studied using scanning electron microscopy. The results showed that there are seven categories and 12 types of sensilla in C. elysii adults: three types of S.Ch, three types of S.T, two types of S.B and one each of B.B, S.Co, S.Ca and S.Cam. There is no difference between male and female in the types of sensilla. Apart from the significant difference in the number and distribution of S.B.2, S.Co and S.Ca between male dorsal and ventral surfaces, there are no significant differences between male and female antennae. In general, the number of sensilla in females is larger than that in males. The results provide a basic reference for future ultrastructure, electrophysiological, and comparative behavioral studies of Carabus species.     

Sensilla on the mouthparts and antennae of the elephant louse,Haematomyzus elephantis piaget (Phthiraptera: Haematomyzidae)

The labial palpus of the elephant louse Haematomyzus elephantis has six sensilla that represent three different types: trichoid, basiconic, and styloconic. Two rows of basiconic sensilla are situated on the dorsal and ventral surfaces of the rostrum, and each row consists of three sensilla. Male and female antennae have 15–17 trichoid sensilla situated on the scape, pedicel, and three antennal annuli. Both sexes have two sensilla basiconica on the dorsal surface of the pedicel near the junction of the scape and pedicel. Two coeloconic (tuft) sensilla are situated on the antennae of both sexes, one sensillum on each of the last two annuli. There are three plate organs, two on the last annulus and one on the penultimate annulus of the male and female antennae. Sexual dimorphism is exhibited in the male and female antennae, in that the male has about twice as many sensilla basiconica on the apex of the last annulus as does the female. The total number of sensilla basiconica on the apex of the male antennae is at least two times the number that is known to be present in any other species of lice. © 1992 Wiley-Liss, Inc.     

Morphology and phylogeny of Henneguya jocu n. sp. (Myxosporea,Myxobolidae), infecting the gills of the marine fish Lutjanus jocu

Henneguya jocu n. sp. (Myxosporea, Myxobolidae) is described from the gill lamellae of the marine teleost fish Lutjanus jocu, with a focus on ultrastructural and molecular features. This myxosporean forms subspherical cysts up to ∼260 μm × 130 μm long, and develops asynchronously. Mature myxospores ellipsoidal with a bifurcated caudal process. Myxospore length 10.9 ± 0.4 μm (n = 50); width, 8.2 ± 0.3 μm (n = 50); and thickness, 2.9 ± 0.5 μm (n = 50). Two equal caudal processes, 34.1 ± 1.0 μm long (n = 50); and total myxospore length, 45.2 ± 1.0 μm (n = 50). Two symmetric valves surround two ellipsoidal polar capsules, 5.0 ± 0.3 × 1.4 ± 0.2 μm (n = 20), each containing an isofilar polar filament forming 4–5 coils along the inner wall of these structures, as well as a binucleated sporoplasm presenting a spherical vacuole and several globular sporoplasmosomes. Both the morphological data and molecular analysis of the SSU rDNA gene identify this parasite as a new species of the genus Henneguya. Maximum Likelihood and Maximum Parsimony analyses further indicate that the parasite clusters within others marine Myxobolidae species, forming a group alongside other Henneguya species described from marine hosts.     

Antennal sensilla on cave species of Australian Paratemnopteryx cockroaches (Blattaria : Blattellidae)

The external morphology and distribution of antennal sensilla of cave-dwelling Australian cockroaches, Paratemnopteryx stonei (Races B and C), P. howarthi and P. sp. nov. (Blattaria : Blattellidae), are described using scanning electron microscopy. Eight major types of sensilla were found. Long and medium-length sensilla chaetica are deeply grooved mechano and contact chemo-receptors with a terminal pore; the long type forms 5–11% and the medium-length 7–22% of all sensilla. Sensilla trichodea type 1 are very slender, non-porous, and form 43–60% of all sensilla. Sensilla trichodea type 2 are stouter, shorter and have wall pores; they form 5–14% of the sensilla. Sensilla basiconica type B are very short, non-porous, inflexible-socket receptors that are known to be hygro- and thermo-receptors; they comprise less than 0.6% of the sensilla. Sensilla basiconica type Gl are short, grooved and have a terminal pore; they form 5–11% of all sensilla. Type G2 are longer with indications of a terminal pore and form 7–10% of the sensilla. Sensilla basiconica type P are short with wall pores, and they form 3–9% of the sensilla. Total sensillar numbers ranged from 5700–8900 for P. stonei, depending on the race and sex, 6950–9950 for P. sp. nov. and 11,700-15,100 for the smaller and possibly more epigean-related P. howarthi. Females had 700–3150 fewer sensilla than males. Comparisons are made between Paratemnopteryx and common epigean cockroaches in relation to sensillar types and numbers.     

Distribution of olfactory and some other antennal sensilla in the male click beetle Agriotes obscurus L. (Coleoptera : Elateridae)

The distribution of 5 types of sensilla was statistically analysed on the 4–10th antennal segments of the male click beetle Agriotes obscurus (Coleoptera : Elateridae). The distribution pattern of the trichoid pheromone receptors (T₂ sensilla) and the olfactory basiconic B₁B₂ sensilla on the antennae of male A. obscurus differs significantly from the distribution pattern of the contact chemoreceptors (T₁ sensilla) and probably the non-olfactory B₇ and D sensilla. A significant peculiarity of the distribution of olfactory sensilla is their location on the antennal segments as 2 separate (dorsal and ventral) fields of sensilla. The numbers of T₂ and B₁B₂ sensilla on dorsal fields of sensilla of the 4–10th segments increase towards the apex of the antenna nearly linearly. On ventral fields of sensilla of the 4–10th antennal segments, the number of B₁B₂ sensilla is nearly uniform; the number of T₂ sensilla in the proximal part of the antenna increases towards the apex, but on distal segments of the antenna their number stabilizes. It is characteristic of both the T₂ and to B₁B₂ sensilla that their numbers are slightly greater on anterior than posterior sides of dorsal sensillar fields, and also greater on posterior than anterior sides of ventral sensillar fields of all antennal segments investigated. We assume that the number of olfactory sensilla on the antennae of male beetles coincides with the distribution of strength of olfactory signal on the antennae of beetles orientating in an odour plume. The distribution patterns of T₂ and B₁B₂ sensilla of the male A. obscurus can be related to some behavioural peculiarities of olfactory orientation (walking or flying and vibrating of the antennae).     

Gall midge olfaction: Pheromone sensitive olfactory neurons in Contarinia nasturtii and Mayetiola destructor

This study describes the morphology and function of the antennal sensilla in two gall midge species, Contarinia nasturtii and Mayetiola destructor, where multi-component sex pheromones have been identified. Both species possess sensilla trichodea, s. coeloconica, s. chaetica and s. circumfila. Sensilla circumfila, which consist of several sensilla that bifurcate and fuse into one structure, are unique for the gall midges. In C. nasturtii s. circumfila are sexually dimorphic. In males, they form elongated loops suspended on cuticular spines, whereas in females they run like worm-like structures directly on the antennal surface. Single sensillum recordings demonstrated that olfactory sensory neurons housed in male s. circumfila in C. nasturtii responded to the female sex pheromone. In M. destructor, s. circumfila were attached to the antennal surface in both sexes, and displayed no response to sex pheromone components.A sexual dimorphism was also found in the number of s. trichodea per antennal segment in both C. nasturtii (male 1 vs. female 7) and M. destructor (male 13 vs. female 10). OSNs located in male M. destructor s. trichodea responded to the sex pheromone. This is the first gall midge single sensillum study, and the first demonstration of the functional significance of s. circumfila.     

Mechanical difference between white and gray matter in the rat cerebellum measured by scanning force microscopy

The mechanical properties of tissues are increasingly recognized as important cues for cell physiology and pathology. Nevertheless, there is a sparsity of quantitative, high-resolution data on mechanical properties of specific tissues. This is especially true for the central nervous system (CNS), which poses particular difficulties in terms of preparation and measurement. We have prepared thin slices of brain tissue suited for indentation measurements on the micrometer scale in a near-native state. Using a scanning force microscope with a spherical indenter of radius ～20 μm we have mapped the effective elastic modulus of rat cerebellum with a spatial resolution of 100 μm. We found significant differences between white and gray matter, having effective elastic moduli of K=294±74 and 454±53 Pa, respectively, at 3 μm indentation depth (n_g=245, n_w=150 in four animals, p<0.05; errors are SD). In contrast to many other measurements on larger length scales, our results were constant for indentation depths of 2–4 μm indicating a regime of linear effective elastic modulus. These data, assessed with a direct mechanical measurement, provide reliable high-resolution information and serve as a quantitative basis for further neuromechanical investigations on the mechanical properties of developing, adult and damaged CNS tissue.