Temperature Adaptation of Active Sodium-Potassium Transport and of Passive Permeability in Erythrocytes of Ground Squirrels

Unidirectional active and passive fluxes of ⁴²K and ²⁴Na were measured in red blood cells of ground squirrels (hibernators) and guinea pigs (nonhibernators). As temperature is lowered, "active" (ouabain-sensitive) K influx and Na efflux were more greatly diminished in guinea pig cells than in those of ground squirrels. The fraction of total K influx which is ouabain sensitive in red blood cells of ground squirrels was virtually constant at all temperatures, whereas it decreased abruptly in guinea pig cells as temperature was lowered. All the passive fluxes (i.e., Na influx, K efflux, and ouabain-insensitive K influx and Na efflux) decreased logarithmically with decrease in temperature in both species, but in ground squirrels the temperature dependence (Q₁₀ 2.5–3.0) was greater than in guinea pig (Q₁₀ 1.6–1.9). Thus, red blood cells of ground squirrel are able to resist loss of K and gain of Na at low temperature both because of relatively greater Na-K transport (than in cells of nonhibernators) and because of reduced passive leakage of ions.     

Immunological relationships of an endogenous guinea pig retrovirus with prototype mammalian type B and type D retroviruses.

A retrovirus endogenous to guinea pig cells was earlier shown to be morphologically similar to type B and type D prototype retroviruses. Molecular hybridization techniques were used to show that guinea pig virus nucleotide sequences are endogenous to both domestic (Cavia porcellus) and indigenous (Cavia aperea) guinea pigs, but cannot be detected in the DNA of either other hystricomorph rodents or other mammals tested. Using radioimmunological techniques designed to detect interspecies relationships, the major internal polypeptide of guinea pig virus (p26) was shown to share three different sets of interspecies antigenic determinants with squirrel monkey retrovirus, viper retrovirus, and mouse mammary tumor virus. Thus, guinea pig virus appears to provide an evolutionary link between type B and D retroviruses.     

Comparative analysis of morphofunctional features of cortical neurons in ground squirrels and guinea pigs under hypothermia

Activity of sensorimotor cortical neurons in the ground squirrel was studied on slices under cooling the incubation medium from 32–34 to 21–26°С. Hypothermia evoked spontaneous firing activity in “silent” neurons and a slight decrease in firing in high-frequency neurons. Changes in the firing rate arose below 27°С and were accompanied by a fall in the spike amplitude. The intensity of hypothermic and post-hypothermic changes in ground squirrels was lower than in guinea pig sensorimotor cortical neurons recorded under the same conditions. In ground squirrels, most hypothermia-resistant were high-frequency (more than 8 spikes/s) neurons, which accounted for 45% of the recorded, while in guinea pigs high-frequency neurons occurred only in 15% of records. By the diameter of cell bodies, the population of sensorimotor cortical neurons was more homogeneous in ground squirrels than in guinea pigs. It is suggested that specific hypothermic changes in sensorimotor cortical neurons of ground squirrels relate to a lower density of K⁺ channels in their plasma membranes, because in the mammalian nervous system the latter open below 27°С due to thermal limitations of the M-cholinergic reaction which blocks these channels.     

Kinetics of the sodium pump in red cells of different temperature sensitivity

Ouabain-sensitive K influx into ground squirrel and guinea pig red cells was measured at 5 and 37 degrees C as a function of external K and internal Na. In both species the external K affinity increases on cooling, being three- and fivefold higher in guinea pig and ground squirrel, respectively, at 5 than at 37 degrees C. Internal Na affinity also increased on cooling, by about the same extent. The effect of internal Na on ouabain-sensitive K influx in guinea pig cells fits a cubic Michaelis-Menten-type equation, but in ground squirrel cells this was true only at high [Na]i. There was still significant ouabain-sensitive K influx at low [Na]i. Ouabain-binding experiments indicated around 800 sites/cell for guinea pig and Columbian ground squirrel erythrocytes, and 280 sites/cell for thirteen-lined ground squirrel cells. There was no significant difference in ouabain bound per cell at 37 and 5 degrees C. Calculated turnover numbers for Columbian and thirteen-lined ground squirrel and guinea pig red cell sodium pumps at 37 degrees C were about equal, being 77-100 and 100-129 s-1, respectively. At 5 degrees C red cells from ground squirrels performed significantly better, the turnover numbers being 1.0-2.3 s-1 compared with 0.42-0.47 s-1 for erythrocytes of guinea pig. The results do not accord with a hypothesis that cold-sensitive Na pumps are blocked in one predominant form.     

Resistance of Erythrocytes of Hibernating Mammals to Loss of Potassium during Hibernation and during Cold Storage

In two species of hibernators, hamsters and ground squirrels, erythrocytes were collected by heart puncture and the K content of the cells of hibernating individuals was compared with that of awake individuals. The K concentration of hamsters did not decline significantly during each bout of hibernation (maximum period of 5 days) but in long-term bouts in ground squirrels (i.e. more than 5 days) the K concentration of cells dropped significantly. When ground squirrels were allowed to rewarm the K content of cells rose toward normal values within a few hours. Erythrocytes of both hamsters and ground squirrels lose K more slowly than those of guinea pigs (nonhibernators) when stored in vitro for up to 10 days at 5°C. In ground squirrels the rate of loss of K during storage is the same as in vivo during hibernation, and stored cells taken from hibernating ground squirrels also lose K at the same rate. The rate of loss of K from guinea pig cells corresponded with that predicted from passive diffusion unopposed by transport. The actual rate of loss of K from ground squirrel cells was slower than such a predicted rate but corresponded with it when glucose was omitted from the storage medium or ouabain was added to it. Despite the slight loss of K that may occur in hibernation, therefore, the cells of hibernators are more cold adapted than those of a nonhibernating mammal, and this adaptation depends in part upon active transport.     

Evolutionary novelty in a rat with no molars

Rodents are important ecological components of virtually every terrestrial ecosystem. Their success is a result of their gnawing incisors, battery of grinding molars and diastema that spatially and functionally separates the incisors from the molars. Until now these traits defined all rodents. Here, we describe a new species and genus of shrew-rat from Sulawesi Island, Indonesia that is distinguished from all other rodents by the absence of cheek teeth. Moreover, rather than gnawing incisors, this animal has bicuspid upper incisors, also unique among the more than 2200 species of rodents. Stomach contents from a single specimen suggest that the species consumes only earthworms. We posit that by specializing on soft-bodied prey, this species has had no need to process food by chewing, allowing its dentition to evolve for the sole purpose of procuring food. Thus, the removal of functional constraints, often considered a source of evolutionary innovations, may also lead to the loss of the very same traits that fuelled evolutionary diversification in the past.     

Diet,bite force and skull morphology in the generalist rodent morphotype

For many vertebrate species, bite force plays an important functional role. Ecological characteristics of a species' niche, such as diet, are often associated with bite force. Previous evidence suggests a biomechanical trade‐off between rodents specialized for gnawing, which feed mainly on seeds, and those specialized for chewing, which feed mainly on green vegetation. We tested the hypothesis that gnawers are stronger biters than chewers. We estimated bite force and measured skull and mandible shape and size in 63 genera of a major rodent radiation (the myomorph sigmodontines). Analysis of the influence of diet on bite force and morphology was made in a comparative framework. We then used phylogenetic path analysis to uncover the most probable causal relationships linking diet and bite force. Both granivores (gnawers) and herbivores (chewers) have a similar high bite force, leading us to reject the initial hypothesis. Path analysis reveals that bite force is more likely influenced by diet than the reverse causality. The absence of a trade‐off between herbivores and granivores may be associated with the generalist nature of the myomorph condition seen in sigmodontine rodents. Both gnawing and chewing sigmodontines exhibit similar, intermediate phenotypes, at least compared to extreme gnawers (squirrels) and chewers (chinchillas). Only insectivorous rodents appear to be moving towards a different direction in the shape space, through some notable changes in morphology. In terms of diet, natural selection alters bite force through changes in size and shape, indicating that organisms adjust their bite force in tandem with changes in food items.     

Differential contractile response of normal vas deferens of rodents in correlation to their calcium and electrolyte levels

The contractile pattern of the vas deferens in three different rodents, rat, guinea pig and mouse was studied in response to adrenaline and noradrenaline. The left vas deferens of rat was more responsive to the graded doses of adrenaline and noradrenaline than the right. The same was also true for guinea pig and mouse vas deferens. This differential response has been correlated with the greater concentrations of calcium and sodium in the right vas deferens in rats and guinea pigs and it might also be related to the levels of membrane-bound and intracellular calmodulin-bound calcium. It is suggested that the left vas deferens might possess more calmodulin-bound calcium than the right, which might have instead, more membrane-bound calcium.     

Bark-stripping by Grey squirrels (Sciurus carolinensis)

To investigate why Grey squirrels strip bark from young trees, squirrel populations and tree quality were studied at 30 English Midland woods. In young beech and sycamore woodlands, the area of bark which Grey squirrels stripped from each tree was strongly related to the phloem width (volume per unit area) of the tree, and not to the phloem sugar content. Variation in bark-stripping between woods was strongly related to average phloem widths in each plantation. The extent of bark-stripping also correlated with juvenile squirrel density, but only in years with relatively little squirrel breeding. There was a tendency for damage to recur in previously damaged areas, in a pattern suggesting that squirrels had learned the habit. The results indicate that bark-stripping can be initiated by young squirrels, perhaps through agonistic gnawing or exploratory feeding, or by older squirrels which have learned the habit, but that severe damage occurs only where the tree phloem is suitable.     

Electromyography and mechanics of mastication in the albino rat.

The masticatory apparatus in the albino rat was studied by means of electromyography and subsequent estimation of muscular forces. The activity patterns of the trigeminal and suprahyoid musculature and the mandibular movements were recorded simultaneously during feeding. The relative forces of the individual muscles in the different stages of chewing cycles and biting were estimated on the basis of their physiological cross sections and their activity levels, as measured from integrated electromyograms. Workinglines and moment arms of these muscles were determined for different jaw positions. In the anteriorly directed masticatory grinding stroke the resultants of the muscle forces at each side are identical; they direct anteriorly, dorsally and slightly lingually and pass along the lateral side of the second molar. Almost the entire muscular resultant force is transmitted to the molars while the temporo-mandibular joint remains unloaded. A small transverse force, produced by the tense symphyseal cruciate ligaments balances the couple of muscle resultant and molar reaction force in the transverse plane. After each grinding stroke the mandible is repositioned for the next stroke by the overlapping actions of three muscle groups: the pterygoids and suprahyoids produce depression and forward shift, the suprahyoids and temporal backward shift and elevation of the mandible while the subsequent co-operation of the temporal and masseter causes final closure of the mouth and starting of the forward grinding movement. All muscles act in a bilaterally symmetrical fashion. The pterygoids contract more strongly, the masseter more weakly during biting than during chewing. The wide gape shifts the resultant of the muscle forces more vertically and moreposteriorly. The joint then becomes strongly loaded because the reaction forces are applied far anteriorly on the incisors. The charateristic angle between the almost horizontal biting force and the surface of the food pellet indicates that the lower incisors produce a chisel-like action. Tooth structure reflects chewing and biting forces. The transverse molar lamellae lie about parallel to the chewing forces whereas perpendicular loading of the occlusal surfaces is achieved by their inclination in the transverse plane. The incisors are loaded approximately parallel to their longitudinal axis, placement that avoids bending forces during biting. It is suggested that a predominantly protrusive musculature favors the effective force transmission to the lower incisors, required for gnawing. By grinding food across transversely oriented molar ridges the protrusive components of the muscles would be utilized best. From the relative weights of the masticatory muscles in their topographical relations with joints, molars and incisors it may be concluded that the masticatory apparatus is a construction adapted to optimal transmission of force from muscles to teeth.     

Donning your enemy's cloak: ground squirrels exploit rattlesnake scent to reduce predation risk

Ground squirrels (Spermophilus spp.) have evolved a battery of defences against the rattlesnakes (Crotalus spp.) that have preyed on them for millions of years. The distinctive behavioural reactions by these squirrels to rattlesnakes have recently been shown to include self-application of rattlesnake scent-squirrels apply scent by vigorously licking their fur after chewing on shed rattlesnake skins. Here, we present evidence that this behaviour is a novel antipredator defence founded on exploitation of a foreign scent. We tested three functional hypotheses for snake scent application--antipredator, conspecific deterrence and ectoparasite defence--by examining reactions to rattlesnake scent by rattlesnakes, ground squirrels and ectoparasites (fleas). Rattlesnakes were more attracted to ground squirrel scent than to ground squirrel scent mixed with rattlesnake scent or rattlesnake scent alone. However, ground squirrel behaviour and flea host choice were not affected by rattlesnake scent. Thus, ground squirrels can reduce the risk of rattlesnake predation by applying rattlesnake scent to their bodies, potentially as a form of olfactory camouflage. Opportunistic exploitation of heterospecific scents may be widespread; many species self-apply foreign odours, but few such cases have been demonstrated to serve in antipredator defence.     

Mechanical advantage of area of origin for the external pterygoid muscle in two murid rodents, Apodemus speciosus and Clethrionomys rufocanus

The actions of masticatory muscles in relation to transverse grinding, associated with forward masticatory movement of the mandible, were investigated by using a mechanical model in the two murid rodents, the Japanese field mouse (Apodemus speciosus: subfamily Murinae) and the gray red-backed vole (Clethrionomys rufocanus: subfamily Arvicolinae). Furthermore, statics of the masticatory system on a sagittal plane while chewing is taking place were also analyzed in these rodents. The inward grinding movements of hemimandibles are generated by the posterior temporalis and internal and external pterygoids in both species. In addition to these muscles, the anterior temporalis also moves the hemimandibles lingually in Apodemus speciosus. The area of origin of the external pterygoid seems more advantageous for transverse grinding in A. speciosus than in Clethrionomys rufocanus. On the basis of the static analysis, the anterodorsal area of origin of the external pterygoid to the upper second and third molars in Clethrionomys rufocanus appears to be an adaptive character to prevent the jaw joints from dislocation during occlusion at a posterior point on the elongated row of cheek teeth.     

The molecular taxonomy and evolution of the guinea pig.

On the basis of 18 protein sequences totaling 2,413 aligned amino acid sites, it is suggested that the guinea pigs and the myomorphs (rat-like rodents) are not monophyletic. Rather, the evolutionary lineage leading to the guinea pig seems to have branched off prior to the divergence among myomorphs, lagomorphs, primates, chiropterans, artiodactyls, and carnivores. It is suggested therefore that the Caviomorpha (guinea pig-like rodents) and possibly the Hystricomorpha (porcupine-like rodents) should be elevated in taxonomic rank and conferred an ordinal status distinct from the Rodentia. This suggestion calls for a reevaluation of the morphological evolution of guinea pigs and further molecular studies on the possibility of paraphyly of the order Rodentia. If the monophyly of rodents holds, it must be concluded that the pattern of molecular evolution in many guinea pig genes has been extremely unusual and that the causes for this pattern should be sought. It is also suggested that claims of large differences in the rate of molecular evolution between guinea pigs and myomorphs may have been exaggerated in many cases as a result of an erroneous phylogenetic position for the guinea pig. The average rate of amino acid replacement in the guinea pig seems to be comparable to that in the rat and the mouse. However, the data indicate that myomorph and caviomorph genes evolve, on average, about two times faster than their human counterparts. Finally, our analysis provides evidence against the hypothesis that the gundi (an African rodent) represents the most ancient rodent lineage.     

A First Generation Comparative Chromosome Map between Guinea Pig (Cavia porcellus) and Humans

The domesticated guinea pig, Cavia porcellus (Hystricomorpha, Rodentia), is an important laboratory species and a model for a number of human diseases. Nevertheless, genomic tools for this species are lacking; even its karyotype is poorly characterized. The guinea pig belongs to Hystricomorpha, a widespread and important group of rodents; so far the chromosomes of guinea pigs have not been compared with that of other hystricomorph species or with any other mammals. We generated full sets of chromosome-specific painting probes for the guinea pig by flow sorting and microdissection, and for the first time, mapped the chromosomal homologies between guinea pig and human by reciprocal chromosome painting. Our data demonstrate that the guinea pig karyotype has undergone extensive rearrangements: 78 synteny-conserved human autosomal segments were delimited in the guinea pig genome. The high rate of genome evolution in the guinea pig may explain why the HSA7/16 and HSA16/19 associations presumed ancestral for eutherians and the three syntenic associations (HSA1/10, 3/19, and 9/11) considered ancestral for rodents were not found in C. porcellus. The comparative chromosome map presented here is a starting point for further development of physical and genetic maps of the guinea pig as well as an aid for genome assembly assignment to specific chromosomes. Furthermore, the comparative mapping will allow a transfer of gene map data from other species. The probes developed here provide a genomic toolkit, which will make the guinea pig a key species to unravel the evolutionary biology of the Hystricomorph rodents.     

Beta-hydroxybutyrate and response to hypoxia in the ground squirrel, Spermophilus tridecimlineatus

1. Previous studies have suggested that elevated ketone levels are associated with increased survival time in rodents exposed to hypoxia. In this study the association between whole blood BHB (beta-hydroxybutyrate) and hypoxic survival time was investigated in hibernating and non-hibernating ground squirrels and in rats. 2. Non-hibernating ground squirrels and rats were exposed to hypoxia (4.5% O2). One hundred per cent of ground squirrels survived 1 hr of hypoxia vs 20% of rats. 3. Ketone levels were significantly higher in ground squirrels than rats during hypoxia, and rats surviving the longest had the highest ketone levels. 4. When hibernation was induced in ground squirrels there was a significant increase in beta-hydroxy-butyrate from 0.45 to 1.6 mM (P = 0.0005). 5. Ground squirrel heart mitochondrial respiratory control ratios and ATP synthesis rates indicated no preferential ketone utilization which might suggest a possible extramitochondrial role of BHB during hypoxia. 6. We conclude that elevated blood BHB levels are associated with increased hypoxic survival and they may have evolved in response to life-threatening hypoxia as experienced during hibernation.     

Cultured cells from renal cortex of hibernators and nonhibernators. Regulation of cell K+ at low temperature

Cells were grown as primary monolayer cultures from kidney cortex of guinea pigs (nonhibernators), hamsters and ground squirrels (both hibernating species). When plates of cells were placed at 5 °C, cells of guinea pigs lost 37% of their K⁺ in 2 h and those of the hibernator lost about 10%.Uptake of ⁴²K into the cells exhibited a simple, single exponential time course at both temperatures. Unidirectional efflux of K⁺ was equal to K⁺ influx in all cultures at 37 °C and, within limits of error, in hibernator cells at 5 °C. Efflux was 3- to 5-fold greater than influx in guinea pig cells at 5 °C.After 2 h in the cold the ouabain-sensitive K⁺ influx remaining (7–15% of that at 37 °C) was about the same in the cells of the 3 species. Cells from active hamsters and from hibernating ground squirrels, however, exhibited significantly greater pump activity after 45 min in the cold (19 and 14%, respectively). The stimulation of K⁺ influx by increasing [K⁺]_o did not show an increase in K_m⁺ at 5 °C in cells of guinea pigs and ground squirrels. Lowering [K⁺]_c and/or raising [Na⁺]_c by treatment in low- and high-K⁺ media caused only slight stimulation of K⁺ influx, except in cells of ground squirrels at 5 °C in which the stimulation was at least 11-times greater than at 37 °C or in cells of guinea pigs at either temperature.This altered kinetic response of K⁺ transport to cytoplasmic ion stimulation with cooling accounted for about one-third of the improved regulation of K⁺ at 5 °C in ground squirrel cells; the other two-thirds was attributable to a greater decrease in K⁺ leak with cooling. The inhibition of active transport by cold in all 3 species was much less severe than that previously seen in any (Na⁺ + K⁺)-ATPase of mammalian cells.     

Comparative physiology of rodent pulmonary macrophages: in vitro functional responses

Since toxicological testing of inhaled materials frequently requires utilization of several species, we have investigated pulmonary macrophage (PM) functional responses and compared the rat model with other rodents. Two strains of rats, three strains of mice, and one strain each of hamster and guinea pig were used in this study. The numbers of recovered cells by bronchoalveolar lavage generally correlated with animal body weight. The one exception was the Syrian Golden hamster from which increased numbers of macrophages were recovered. Cellular differential data obtained from lavaged cytocentrifuge preparations demonstrated that PM's account for greater than 97% of recoverable free lung cells for all species except the guinea pig, which contains a resident population of eosinophils. Cell morphology studies indicated that hamster PM exhibited the highest proportion of ruffled PM and demonstrated the highest phagocytic activity, whereas mouse PM phagocytic activity was significantly reduced compared with the other three species. In addition, chemotaxis studies showed that rat PM migrated best to zymosan-activated, complement-dependent chemoattractants, whereas hamster PM demonstrated an enhanced chemotactic response to N-formyl peptides. The results of these studies suggest that the rat may be the most efficient species for clearing inhaled particles, whereas hamsters and guinea pigs may best respond to bacteria.     

Biological characterization of acute infection with ground squirrel hepatitis virus.

Ground squirrel hepatitis virus (GSHV) is a small DNA virus, structurally and antigenically related to the human hepatitis B virus, which occurs naturally among certain wild populations of ground squirrels (P. L. Marion et al., Proc. Natl. Acad. Sci. U.S.A. 77:2941-2945, 1980). Serum from naturally infected animals was used to transmit GSHV in the laboratory by parenteral inoculation of susceptible squirrels. Sixty percent of recipient animals developed viral surface antigenemia after a latent period of 2 to 3 months; three of these animals have remained viremic for over 9 months. Like hepatitis B virus, GSHV demonstrates marked hepatotropism, with viral DNA detected in significant quantities only in the liver, where an average of 6 X 10(2) to 6 X 10(3) viral DNA molecules per cell were found by molecular hybridization. However, histological signs of liver injury after acute infection are minimal. In contrast to infection of its natural host, parenteral administration of GSHV to rats, mice, guinea pigs, and hamsters did not result in demonstrable antigenemia, suggesting that the host range of GSHV, like that of hepatitis B virus, is narrow.     

Occlusal Pattern in Paulchoffatiid Multituberculates and the Evolution of Cusp Morphology in Mammaliamorphs with Rodent-like Dentitions

Multituberculates developed a very complex masticatory apparatus during their long evolutionary history from the Jurassic to the Paleogene. Besides their rodent-like elongated incisors and diastemata, Cenozoic cimolodont Multituberculata display masticatory movements involving two distinct cycles in the mastication. An orthal slicing-crushing cycle associated with an enlarged lower fourth premolar precedes a palinal grinding cycle linked to upper molars with three longitudinal rows of cusps. With their plesiomorphic lower premolars and upper molars, the Late Jurassic/Early Cretaceous multituberculate family Paulchoffatiidae can provide the key for the understanding of the origin of the complex mastication of the Cimolodonta. Using for the first time propagation phase contrast Synchrotron X-Ray microtomography to perform both microwear and topographic analyses in order to characterize the mastication of Paulchoffatiidae, we digitized dental material from the Late Jurassic of the Guimarota Coal Mine (Leiria, Portugal) at the European Synchrotron Facility (Grenoble, France). Mastication in Paulchoffatiidae is characterized by a palinal grinding cycle. In contrast to Cimolodonta, no evidence of an orthal slicing-crushing cycle has been observed: the lower premolars mainly have a grinding function like the molars as they do exhibit buccal attrition facets bearing longitudinal striations. Nevertheless, the slightly oblique striations observed on the mesial part of the paulchoffatiid lower premolars possibly presage the orthal phase of the Cimolodonta. Our topographic analysis indicates that a strong relationship between individual cusp shape and direction of chewing is emphasized in rodents and rodent-like Mammaliamorpha such as Cimolodonta and Tritylodonta. Surprisingly, this relationship is not evident in Paulchoffatiidae. This unexpected result can be explained by the non-involvement in the attrition of many premolar cusps in Paulchoffatiidae, as indicated by our microwear analysis. The stronger the attrition, the more the direction of the masticatory movements influences the cusp morphology in Mammaliamorpha.     

Glia–Neuron Interactions in the Sensory-Motor Cortex of Warm-Blooded Animals (Guinea Pigs and Ground Squirrels) with Different Habitat Conditions and the M-Cholinergic Reaction of the Brain

The glia–neuron interactions were analyzed in the sensory-motor cortex of guinea pigs and ground squirrels (Spermophilus undulatus) during the active summer months. The glial cells were more concentrated in close proximity (15–25 μm) to neurons (38% in guinea pigs and 22.4% in ground squirrels). A more concentrated distribution of glial cells might be very necessary for spontaneous inactive nerve cells (37.2% in guinea pigs and 23% in ground squirrels), since these neurons are associated with the highest energy demand during their functioning and are most susceptible to disturbances of ion homeostasis. The network structure of glia and the close contact between glial cells and brain capillaries provide additional energy for neurons and stabilize the ion balance in the extracellular medium. Glial density in the sensory-motor cortex of ground squirrels is 3 times higher than that in the cortex of guinea pigs. The high content of glial cells in the ground-squirrel cortex is the most important protective factor for survival of animals during long-term hibernation, when the diffusion of K⁺ ions from nerve cells drastically increases due to the high temperature sensitivity of the M-cholinergic response.