Balancing function of the masticatory muscles during incisal biting in two murid rodents,Apodemus speciosus and Clethrionomys rufocanus

The functional significance of masticatory muscle direction was estimated using a mechanical model in two murid rodents: the Japanese field mouse (Apodemus speciosus) and the gray red-backed vole (Clethrionomys rufocanus). Theoretical analyses of the data suggest that a balancing mechanism among the muscle forces occurs during incisal power stroke. The activation of the large deep masseter in both murids results in marked tensile separation of two hemimandibles at the flexible mandibular symphysis. Activation of the internal pterygoid decreases this large tensile force at the symphysis more efficiently than other muscles. The lines of action of the deep masseter and internal pterygoid are aligned to produce such a balancing function in both species studied here. The resultant force generated by the deep masseter on both sides is opposite in direction to the reaction force at the lower incisor tip. Therefore, the large deep masseter forms an effective mandibular support mechanism when the reaction forces during biting push the mandible downward. Because of the area of insertion and the line of action, the posterior temporalis appears to have an important role in stabilizing the position of the mandibular condyle in the glenoid fossa during incisal biting. J. Morphol. 236:49–56, 1998. © 1998 Wiley-Liss, Inc.     

Jaw muscle functional anatomy in northern grasshopper mouse, Onychomys leucogaster, a carnivorous murid

The jaw muscle anatomy of the northern grasshopper mouse, Onychomys leucogaster, was observed and the mechanical basis of the insectivorous/carnivorous adaptations were examined. Compared with Peromyscus maniculatus, a granivorous relative of Onychomys, there is a reduction of some aponeuroses within the masseter deep layer. This characteristic indicates that shearing meat or crushing arthropod exoskeletons requires less occlusal pressure than does grinding plant material. In Onychomys both the anterior and posterior portions of the masseter deep layer are more anterodorsally inclined, so that the line of action of the masseter lies further from the jaw joint than in Peromyscus. A strong incisal bite for killing vertebrates such as other rodents can be produced by a jaw mechanism with the high lever advantage of this muscle, which compensates for the decline in muscle mass. Our quantitative analysis suggests that the disappearance of an aponeurosis along the zygomatic plate in Onychomys decreases the stretch of the corresponding muscle, i.e., the anterior fibers of the masseter deep layer, accompanying jaw opening, and increases the maximum gape necessary for hunting large prey.     

Masticatory Muscle Anatomy and Feeding Efficiency of the American Beaver, <Emphasis Type="Italic">Castor canadensis</Emphasis> (Rodentia,Castoridae)

Beavers are well-known for their ability to fell large trees through gnawing. Yet, despite this impressive behavior, little information exists on their masticatory musculature or the biomechanics of their jaw movements. It was hypothesized that beavers would have a highly efficient arrangement of the masticatory apparatus, and that gnawing efficiency would be maintained at large gape. The head of an American beaver, Castor canadensis, was dissected to reveal the masticatory musculature. Muscle origins and insertions were noted, the muscles were weighed and fiber lengths measured. Physiological cross-sectional areas were determined, and along with the muscle vectors, were used to calculate the length of the muscle moment arms, the maximum incisor bite force, and the proportion of the bite force projected along the long axis of the lower incisor, at occlusion and 30° gape. Compared to other sciuromorph rodents, the American beaver was found to have large superficial masseter and temporalis muscles, but a relatively smaller anterior deep masseter. The incisor bite force calculated for the beaver (550–740 N) was much higher than would be predicted from body mass or incisor dimensions. This is not a result of the mechanical advantage of the muscles, which is lower than most other sciuromorphs, but is likely related to the very high percentage (>96 %) of bite force directed along the lower incisor long axis. The morphology of the skull, mandible and jaw-closing muscles enable the beaver to produce a very effective and efficient bite, which has permitted beavers to become highly successful ecosystem engineers.     

Species differences among various rodents in the conversion of 7alpha-hydroxycholesterol in liver microsomes

Our previous study demonstrated that there are species differences among vertebrates in their conversion of 7alpha-hydroxycholesterol (7HC) to 7-ketocholesterol (7KC). To examine this further, we investigated the differences in the products of 7alpha-hydroxycholesterol in various species of male muroid rodents. Adult male Syrian hamsters (Mesocricetus auratus), dwarf hamsters (Phodopus rovolovskii), Djungarian hamsters (Phodopus sungorus), Chinese hamsters (Cricetulus griseus), rat-like hamsters (Tscherskia triton), and hispid cotton rats (Sigmodon hispidus) were used. Microsomal fractions were prepared from their livers, and the activities of the enzymes that participate in the dehydrogenation of 7alpha-hydroxycholesterol were determined by measuring the products using high-performance liquid chromatography. 7alpha-hydroxycholesterol was converted to both 7alpha-hydroxy-4-cholesten-3-one (7HCO) and 7-ketocholesterol in all of the hamsters tested. However, in the rat-like hamster and the hispid cotton rat, 7alpha-hydroxycholesterol was converted mostly to 7alpha-hydroxy-4-cholesten-3-one, as also observed in the rat (Rattus norvegicus). The results suggest that microsomal enzyme activity in the conversion of 7alpha-hydroxycholesterol to 7-ketocholesterol varies considerably, even within the subfamily Cricetinae and the family Muridae.     

Two novel cricetine mitogenomes: Insight into the mitogenomic characteristics and phylogeny in Cricetinae (Rodentia: Cricetidae)

Both Cricetus cricetus and Phodopus sungorus mitochondrial genomes (mitogenomes) were sequenced and elaborated for the first time in the present study. Their mitogenomes contained 37 genes and showed typical characteristics of the vertebrate mitogenome. Comparative analysis of 10 cricetine mitogenomes indicated that they shared similar characteristics with those of other cricetines in terms of genes arrangement, nucleotide composition, codon usage, tRNA structure, nucleotide skew and the origin of replication of light strand. Phylogenetic relationship of the subfamily Cricetinae was reconstructed using mitogenomes data with the methods of Bayesian Inference and Maximum Likelihood. Phylogenetic analysis indicated that Cricetulus kamensis was at basal position and phylogenetically distant from all other Cricetulus species but had a close relationship with the group of Phodopus, and supported that the genus Urocricetus deserved as a separate genus rank. The phylogenetic status of Tscherskia triton represented a separate clade corresponding to a diversified cricetine lineage (Cricetulus, Allocricetulus, and Cricetus).     

Trends in the Actions of Mammalian Masticatory Muscles

The actions of the masticatory muscles of a variety of mammalsin which feeding behavior and the configuration of the masticatoryapparatus differ have been reported. The most common approachused in these studies involves (1) obtaining a good anatomicalperception of the musculature, (2) deriving a theoretical modelof the actions of these muscles during jaw movement, and (3)testing this model by recording muscle activity and jaw movementssimultaneously. A catalogue of the activity patterns in eleven species of mammalsduring food reduction reveals certain trends in the actionsof the masticatory muscles. Horizontal jaw movements are generatedprimarily by differential activities of the deep temporalis,superficial masseter, and medial pterygoid. Vertical movementsand the maintenance of tooth to food contact apparently areproduced by action of the superficial temporalis, deep masseter,and zygomaticomandibularis. Thus, horizontal movements are seeminglygenerated by muscles having fibers arranged in marked anteroposteriordirection, whereas vertical movements are generated by muscleshaving more or less vertically arranged fibers. The asymmetry of jaw movement and the muscular activity generatingit suggest that mastication involves an interactionbetween anunbalanced and flexible functional unit (muscles) and a balancedand stable structural unit (skull and teeth). Thus, any unbalancingof the structural unit results in a further unbalancing of themasticatory process.     

Form and function of the masticatory musculature in the tree sloths, Bradypus and Choloepus

The tree sloths, Bradypus and Choloepus, show unusual masticatory specializations, compared to each other and to other mammals. Both have an incomplete zygomatic arch with descending jugal process, a complex superficial masseter, a large temporalis and medial pterygoid musculature, and a lateral pterygoid with two heads. In Choloepus the deep masseter and zygomaticomandibularis are typical when compared to other mammals. However, in Bradypus there is an ascending jugal process from which enlarged and vertically oriented deep masseter and zygomaticomandibularis muscles originate. Although both sloths are folivores, the anterior teeth in Choloepus are caniniform, while those of Bradypus have lost such elongation. In both sloths the glenoid cavity is similarly located; however, in Bradypus the craniomandibular joint is raised above the occlusal plane, and the pterygoid flanges are elongated. Prediction of the evolutionary sequence of cranial changes from Choloepus-like (primitive) to Bradypus-like (derived) morphology is based upon the most parsimonious model of masseter-medial pterygoid complex changes for masticatory efficiency improvement. The model proposes that the condylar neck in Bradypus was elongated and that this single change predicated a series of other structural changes. Mandibular movement patterns in both sloths showed anteromedially directed unilateral power strokes as in other mammals. Puncture-crushing, tooth-sharpening, and chewing cycles are distinct in Choloepus, less so in Bradypus. The masticatory rate is slow in sloths compared to other mammals of similar body size, averaging 590 ms per cycle for Choloepus and 510 ms for Bradypus.     

Occurrence and distribution of muscle spindles in masticatory and suprahyoid muscles of the rat.

The occurrence and distribution of muscle spindles was studied in histochemically and conventionally stained serial cross sections of 6-week-old and adult rat masticatory and suprahyoid muscles. Spindles were present in moderate to large numbers in jaw closers, but they were absent in jaw openers and two of four muscles of an accessory suprahyoid group. In jaw closers, 67% or more of the total spindle population was concentrated relatively distant from the temporomandibular joint, in muscle portions which contained large numbers of extrafusal fibers reacting strongly for oxidative enzymes. Because of their location, spindles in these portions should be stretched more and, subsequently, should respond with a greater afferent discharge at any given muscle length than spindles situated nearer to the joint. Spindles in jaw closers, especially the medial pterygoid and deep masseter, often occurred in clusters and complex forms near the terminal branching of intramuscular nerve trunks. No such concentrations were seen in the two muscles of the accessory suprahyoid group that had spindles. The association in jaw closers of spindles with extrafusal fibers high in oxidative enzyme activity is consistent with the view that spindles are the sensory component of a reflex system that recruits these fibers for finely-graded contractions in response to small internal length-changes of the muscle (Botterman et al., '78); however, in jaw openers and two muscles of the accessory suprahyoid group, the absence of spindles, coupled with the presence of large populations of extrafusal fibers high in oxidative enzyme activity, is not easily reconciled with this concept.     

Molecular phylogeny of the Cricetinae subfamily based on the mitochondrial cytochrome b and 12S rRNA genes and the nuclear vWF gene

Despite some popularity of hamsters as pets and laboratory animals there is no reliable phylogeny of the subfamily Cricetinae available so far. Contradicting views exist not only about the actual number of species but also concerning the validity of several genera. We used partial DNA sequences of two mitochondrial (cytochrome b, 12S rRNA) and one partial nuclear gene (von Willebrand Factor exon 28) to provide a first gene tree of the Cricetinae based on 15 taxa comprising six genera. According to our data, Palaearctic hamsters fall into three distinct phylogenetic groups: Phodopus, Mesocricetus, and Cricetus-related species which evolved during the late Miocene about 7-12MY ago. Surprisingly, the genus Phodopus, which was previously thought to have appeared during the Pleistocene, forms the oldest clade. The largest number of extant hamster genera is found in a group of Cricetus-related hamsters. The genus Cricetulus itself proved to be not truly monophyletic with Cricetulus migratorius appearing more closely related to Tscherskia, Cricetus, and Allocricetulus. We propose to place the species within a new monotypic genus. Molecular clock calculations are not always in line with the dating of fossil records. DNA based divergence time estimates as well as taxonomic relationships demand a reevaluation of morphological characters previously used to identify fossils and extant hamsters.     

Masticatory motor patterns in ungulates: a quantitative assessment of jaw-muscle coordination in goats, alpacas and horses

We investigated patterns of jaw-muscle coordination during rhythmic mastication in three species of ungulates displaying the marked transverse jaw movements typical of many large mammalian herbivores. In order to quantify consistent motor patterns during chewing, electromyograms were recorded from the superficial masseter, deep masseter, posterior temporalis and medial pterygoid muscles of goats, alpacas and horses. Timing differences between muscle pairs were evaluated in the context of an evolutionary model of jaw-muscle function. In this model, the closing and food reduction phases of mastication are primarily controlled by two distinct muscle groups, triplet I (balancing-side superficial masseter and medial pterygoid and working-side posterior temporalis) and triplet II (working-side superficial masseter and medial pterygoid and balancing-side posterior temporalis), and the asynchronous activity of the working- and balancing-side deep masseters. The three species differ in the extent to which the jaw muscles are coordinated as triplet I and triplet II. Alpacas, and to a lesser extent, goats, exhibit the triplet pattern whereas horses do not. In contrast, all three species show marked asynchrony of the working-side and balancing-side deep masseters, with jaw closing initiated by the working-side muscle and the balancing-side muscle firing much later during closing. However, goats differ from alpacas and horses in the timing of the balancing-side deep masseter relative to the triplet II muscles. This study highlights interspecific differences in the coordination of jaw muscles to influence transverse jaw movements and the production of bite force in herbivorous ungulates.     

Control of jaw movements in two species of macropodines (Macropus eugenii and Macropus rufus).

The masticatory motor patterns of three tammar wallabies and two red kangaroos were determined by analyzing the pattern of electromyographic (EMG) activity of the jaw adductors and correlating it with lower jaw movements, as recorded by digital video and videoradiography. Transverse jaw movements were limited by the width of the upper incisal arcade. Molars engaged in food breakdown during two distinct occlusal phases characterized by abrupt changes in the direction of working-side hemimandible movement. Separate orthal (Phase I) and transverse (Phase II) trajectories were observed. The working-side lower jaw initially was drawn laterally by the balancing-side medial pterygoid and then orthally by overlapping activity in the balancing- and working-side temporalis and the balancing-side superficial masseter and medial pterygoid. Transverse movement occurred principally via the working-side medial pterygoid and superficial masseter. This pattern contrasted to that of placental herbivores, which are known to break down food when they move the working-side lower jaw transversely along a relatively longer linear path without changing direction during the power stroke. The placental trajectory results from overlapping activity in the working- and balancing-side adductor muscles, suggesting that macropods and placental herbivores have modified the primitive masticatory motor pattern in different ways.     

Control of jaw movements in two species of macropodines (Macropus eugenii and Macropus rufus)

The masticatory motor patterns of three tammar wallabies and two red kangaroos were determined by analyzing the pattern of electromyographic (EMG) activity of the jaw adductors and correlating it with lower jaw movements, as recorded by digital video and videoradiography. Transverse jaw movements were limited by the width of the upper incisal arcade. Molars engaged in food breakdown during two distinct occlusal phases characterized by abrupt changes in the direction of working-side hemimandible movement. Separate orthal (Phase I) and transverse (Phase II) trajectories were observed. The working-side lower jaw initially was drawn laterally by the balancing-side medial pterygoid and then orthally by overlapping activity in the balancing- and working-side temporalis and the balancing-side superficial masseter and medial pterygoid. Transverse movement occurred principally via the working-side medial pterygoid and superficial masseter. This pattern contrasted to that of placental herbivores, which are known to break down food when they move the working-side lower jaw transversely along a relatively longer linear path without changing direction during the power stroke. The placental trajectory results from overlapping activity in the working- and balancing-side adductor muscles, suggesting that macropods and placental herbivores have modified the primitive masticatory motor pattern in different ways.     

Jaw-muscle electromyography during chewing in Belanger's treeshrews (Tupaia belangeri)

We examined masseter and temporalis recruitment and firing patterns during chewing in five male Belanger's treeshrews (Tupaia belangeri), using electromyography (EMG). During chewing, the working-side masseters tend to show almost three times more scaled EMG activity than the balancing-side masseters. Similarly, the working-side temporalis muscles have more than twice the scaled EMG activity of the balancing-side temporalis. The relatively higher activity in the working-side muscles suggests that treeshrews recruit less force from their balancing-side muscles during chewing. Most of the jaw-closing muscles in treeshrews can be sorted into an early-firing or late-firing group, based on occurrence of peak activity during the chewing cycle. Specifically, the first group of jaw-closing muscles to reach peak activity consists of the working-side anterior and posterior temporalis and the balancing-side superficial masseter. The balancing-side anterior and posterior temporalis and the working-side superficial masseter peak later in the power stroke. The working-side deep masseter peaks, on average, slightly before the working-side superficial masseter. The balancing-side deep masseter typically peaks early, at about the same time as the balancing-side superficial masseter. Thus, treeshrews are unlike nonhuman anthropoids that peak their working-side deep masseters early and their balancing-side deep masseters late in the power stroke. Because in anthropoids the late firing of the balancing-side deep masseter contributes to wishboning of the symphysis, the treeshrew EMG data suggest that treeshrews do not routinely wishbone their symphyses during chewing. Based on the treeshrew EMG data, we speculate that during chewing, primitive euprimates 1) recruited more force from the working-side jaw-closing muscles as compared to the balancing-side muscles, 2) fired an early group of jaw-closing muscles followed by a second group of muscles that peaked later in the power stroke, 3) did not fire their working-side deep masseter significantly earlier than their working-side superficial masseter, and 4) did not routinely fire their balancing-side deep masseter after the working-side superficial masseter.     

Architecture of the masticatory apparatus in eastern raccoons (Procyon lotor lotor)

The structure and function of the masticatory apparatus of raccoons resemble those found in carnivores. In this study, the architecture of the skull, dentition, and masticatory apparatus is described, and a model is proposed that suggests a mechanism used by raccoons to reduce different foods. The model suggests that jaw movements are similar to those of cats, the posterior regions of the superficial and deep parts of the temporalis and the anterior region of the medial pterygoid generate horizontal jaw movements, and the anterior portions of the superficial and deep temporalis as well as portions of the masseteric complex generate vertical closing movement. The distributions of slow, fast fatigable, and fast fatigue-resistant fibers for the temporalis and masseteric complex are related to the possible actions of these muscles during mastication, as are the regional cross-sectional areas of the masticatory muscles.     

Structure and direction of jaw adductor muscles as herbivorous adaptations in <Emphasis Type="Italic">Neotoma mexicana</Emphasis> (Muridae,Rodentia)

The herbivorous adaptations of the jaw adductor muscles in Neotoma mexicana were clarified by a comparative study with an unspecialized relative, Peromyscus maniculatus. In P. maniculatus, the anterior part of the deep masseter arises entirely from the lateral side of an aponeurosis, i.e., superior zygomatic plate aponeurosis, whereas N. mexicana has an additional aponeurosis for this part of the muscle, and the fibers attach on both sides of the superior zygomatic plate aponeurosis. Although the structure of the temporalis muscle is nearly identical in the two genera, a clear aponeurosis of origin occurs only in N. mexicana. These characteristics allow fibrous tissues to be processed with a large occlusal force. The deep masseter, internal pterygoid, and external pterygoid muscles of N. mexicana incline more anterodorsally than those of P. maniculatus. The transverse force component of these muscles relative to whole muscle force is smaller in N. mexicana than in P. maniculatus, with the exception of the internal pterygoid. The anterior part of the temporalis muscle of N. mexicana is specialized to produce occlusal pressure. These findings suggest that in N. mexicana a large anterior force is required to move the heavy mandible, due to the hypsodont molars, against frictional force from food, and that the posterior pull of the temporalis, which adjusts the forward force by the other jaw adductor muscles to a suitable level, need not be large for the mandibular movement.     

Masseter electromyography during chewing in ring-tailed lemurs (Lemur catta)

We examined masseter recruitment and firing patterns during chewing in four adult ring-tailed lemurs (Lemur catta), using electromyography (EMG). During chewing of tougher foods, the working-side superficial masseter tends to show, on average, 1.7 times more scaled EMG activity than the balancing-side superficial masseter. The working-side deep masseter exhibits, on average, 2.4 times the scaled EMG activity of the balancing-side deep masseter. The relatively larger activity in the working-side muscles suggests that ring-tailed lemurs recruit relatively less force from their balancing-side muscles during chewing. The superficial masseter working-to-balancing-side (W/B) ratio for lemurs overlaps with W/B ratios from anthropoid primates. In contrast, the lemur W/B ratio for the deep masseter is more similar to that of greater galagos, while both are significantly larger than W/B ratios of anthropoids. Because ring-tailed lemurs have unfused and hence presumably weaker symphyses, these data are consistent with the symphyseal fusion-muscle recruitment hypothesis stating that symphyseal fusion in anthropoids provides increased strength for resisting forces created by the balancing-side jaw muscles during chewing. Among the masseter muscles of ring-tailed lemurs, the working-side deep masseter peaks first on average, followed in succession by the balancing-side deep masseter, balancing-side superficial masseter, and finally the working-side superficial masseter. Ring-tailed lemurs are similar to greater galagos in that their balancing-side deep masseter peaks well before their working-side superficial masseter. We see the opposite pattern in anthropoids, where the balancing-side deep masseter peaks, on average, after the working-side superficial masseter. This late activity of the balancing-side deep masseter in anthropoids is linked to lateral-transverse bending, or wishboning, of their mandibular symphyses. Subsequently, the stresses incurred during wishboning are hypothesized to be a proximate reason for strengthening, and hence fusion, of the anthropoid symphysis. Thus, the absence of this muscle-firing pattern in ring-tailed lemurs with their weaker, unfused symphyses provides further correlational support for the symphyseal fusion late-acting balancing-side deep masseter hypothesis linking wishboning and symphyseal strengthening in anthropoids. The early peak activity of the working-side deep masseter in ring-tailed lemurs is unlike galagos and most similar to the pattern seen in macaques and baboons. We hypothesize that this early activity of the working-side deep masseter moves the lower jaw both laterally toward the working side and vertically upward, to position it for the upcoming power stroke. From an evolutionary perspective, the differences in peak firing times for the working-side deep masseter between ring-tailed lemurs and greater galagos indicate that deep masseter firing patterns are not conserved among strepsirrhines.     

An atypical Bronze Age mandible from Żerniki Górne,Poland

The unusual anatomical features of a Bronze Age mandible are described. Of particular interest were the shelf-like ridges which were located approximately midway along the rami. These ridges probably formed the insertion areas for the masseter muscles. Of two possible explanations presented to account for the unusual morphological variations, the more likely is the congenital absence of the superficial portions of the masseter muscles. However, an alternative explanation is presented suggesting the presence of shortened superficial portions of the masseter muscle which could explain the contour of the shelf.     

Loading patterns and jaw movements during mastication in Macaca fascicularis: a bone-strain, electromyographic, and cineradiographic analysis

Rosette strain gage, electromyography (EMG), and cineradiographic techniques were used to analyze loading patterns and jaw movements during mastication in Macaca fascicularis. The cineradiographic data indicate that macaques generally swallow frequently throughout a chewing sequence, and these swallows are intercalated into a chewing cycle towards the end of a power stroke. The bone strain and jaw movement data indicate that during vigorous mastication the transition between fast close and the power stroke is correlated with a sharp increase in masticatory force, and they also show that in most instances the jaws of macaques are maximally loaded prior to maximum intercuspation, i.e. during phase I (buccal phase) occlusal movements. Moreover, these data indicate that loads during phase II (lingual phase) occlusal movements are ordinarily relatively small. The bone strain data also suggest that the duration of unloading of the jaw during the power stroke of mastication is largely a function of the relaxation time of the jaw adductors. This interpretation is based on the finding that the duration from 100% peak strain to 50% peak strain during unloading closely approximates the half-relaxation time of whole adductor jaw muscles of macaques. The EMG data of the masseter and medial pterygoid muscles have important implications for understanding both the biomechanics of the power stroke and the external forces responsible for the "wishboning" effect that takes place along the mandibular symphysis and corpus during the power stroke of mastication. Although both medial pterygoid muscles reach maximum EMG activity during the power stroke, the activity of the working-side medial pterygoid peaks after the balancing-side medial pterygoid. Associated with the simultaneous increase of force of the working-side medial pterygoid and the decrease of force of the balancing-side medial pterygoid is the persistently high level of EMG activity of the balancing-side deep masseter (posterior portion). This pattern is of considerable significance because the direction of force of both the working-side medial pterygoid and the balancing-side deep masseter are well aligned to aid in driving the working-side lower molars across the upper molars in the medial direction during unilateral mastication.(ABSTRACT TRUNCATED AT 400 WORDS)     

Studies on sex chromosomes of four hamster species: Cricetus cricetus, Cricetulus griseus, Mesocricetus auratus, and Phodopus sungorus.

In this paper, we present an analysis of the sex chromosomes of four hamster species after application of different staining techniques. The mitotic X chromosomes show a striking similarity in G-banding pattern but rather great differences in their C-banding patterns. A presumably homologous euchromatic segment that exhibits two distinct G-bands appears in the X chromosome of each species. The Y chromosome of Cricetus cricetus is in contrast to those of the other species, because it reveals a relatively well-differentiated G- and C-banding pattern. In meiotic metaphase I, interstitial chiasmata can be found in the sex bivalents of Cricetus cricetus and Cricetulus griseus, whereas the gonosomes of Mesocricetus auratus and Phodopus sungorus sungorus are terminally associated. The regions that are involved in pairing or association are always heterochromatic.