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Aspects of social life in the fat-tailed dwarf lemur (Cheirogaleus medius): inferences from body weights and trapping data.

Body weight is an important attribute of any animal and influences its biology in many ways. In the case of the fat-tailed dwarf lemur (Cheirogaleus medius), this is of special interest because animals of this species store fat to survive extended torpor phases during the austral winter and, as a result, show extreme seasonal variation in body weight. In this paper, I present field data from a 20-month study at the Forestry Station of Ampijoroa in northwestern Madagascar on body weights of repeatedly trapped individuals combined with trapping results to investigate the general biology and notably the social life of C. medius. The most remarkable physical characteristic of wild C. medius is a dramatic weight-loss in males in September following emergence from torpor. Trapping results showed that adult males emerge from torpor earlier than females and offspring, even though food is still scarce. Early emergence from torpor puts males at an energetic disadvantage and confers no advantage in terms of access to females. Observation data from nightly follows indicate that males patrol their home range, probably to guarantee its integrity for the coming active season and to secure the food supply for their families. The high energetic costs of this strategy is argued to represent a form of paternal investment. Body weight data of repeatedly captured individuals showed that they may reach adult size and sexual maturity at 2 years of age but are not socially mature before the age of 3 years. The data presented may serve as a guide for keeping C. medius at a suitable weight in captivity.

Animals↗

Comparative morphology and histochemistry of glands associated with the vomeronasal organ in humans, mouse lemurs, and voles.

The vomeronasal organ (VNO) is a chemosensory structure of the vertebrate nasal septum that has been recently shown to exist in nearly all adult humans. Although its link to reproductive behaviors has been shown in some primates, its functionality in humans is still debated. Some authors have suggested that the human VNO has the capacity to detect pheromones, while others described it as little more than a glandular pit. However, no studies have utilized histochemical techniques that would reveal whether the human VNO functions as a generalized gland duct or a specialized chemosensory organ. Nasal septal tissue from 13 humans (2-86 years old) were compared to that of two adult lemurs (Microcebus murinus) and eight adult voles (four Microtus pennsylvanicus and four Microtus ochrogaster). Sections at selected intervals of the VNO were stained with periodic acid-Schiff (PAS), alcian blue (AB), AB-PAS, and PAS-hematoxylin procedures. Results revealed typical well-developed VNOs with tubuloacinar glands in Microtus and Microcebus. VNO glands were AB-negative and PAS-positive in voles and mouse lemurs. Homo differed from Microtus and Microcebus in having more branched, AB and PAS-positive glands that emptied into the VNO lumen. Furthermore, the human VNO epithelium had unicellular mucous glands (AB and PAS-positive) and cilia, similar to respiratory epithelia. These results demonstrate unique characteristics of the human VNO which at once differs from glandular ducts (e.g., cilia) and also from the VNOs of mammals possessing demonstrably functional VNO.

Adolescent↗

Species-specific usage of sleeping sites in two sympatric mouse lemur species (Microcebus murinus and M. ravelobensis) in northwestern Madagascar.

We investigated the sleeping site ecology of two sympatric mouse lemur species (Microcebus murinus and M. ravelobensis) in northwestern Madagascar during the second half of the dry season with respect to the type, quality, and usage pattern of the sleeping sites, as well as to social sleeping habits and response to potential threats. The type and quality of the sleeping sites differed between the two species. M. murinus used protected wooden shelters (tree holes) more frequently than M. ravelobensis, and M. ravelobensis used a broader variety of less protected sites (e.g., branches, lianas, and leaves) than M. murinus. Whereas male M. murinus usually slept alone, and female M. murinus mostly slept in groups, both sexes of M. ravelobensis slept in mixed-sex sleeping groups. M. murinus relied on crypsis in their sleeping sites, whereas M. ravelobensis regularly showed a flight response to the approach of an observer. This behavioral difference could indicate an adaptation to a higher predation risk in less protected sleeping sites. Whereas female M. murinus showed a high site fidelity, male M. murinus and both sexes of M. ravelobensis frequently changed their sleeping sites, which may also be interpreted as an antipredator strategy. The results are discussed with respect to three possible ecological explanations: interspecies competition, restricted resource availability, and niche differentiation. The latter is the most likely explanation for these interspecific differences.

Animals↗

Colony specificity in a social call of mouse lemurs (Microcebus ssp.).

Nocturnal primate species are often difficult to discriminate by gross visual bodily characteristics. This is also true for the world's smallest primate taxon, the Malagasy mouse lemurs. Recent findings imply that this taxon contains sibling species that can be diagnosed noninvasively by their species-specific advertisement call. We used comparative bioacoustics in order to explore variation of this call type and to assess species status of three European colonies. Acoustic variation was compared within and between colonies as well as with known species-specific differences. It was further related to morphological and genetic variations to investigate in how far it covaries with them. Results show that acoustic and genetic differences revealed by random amplified polymorphic DNA (RAPD) fingerprinting separated colonies reliably, but were on a different level than known species-specific differences. A Mantel test showed that acoustic differences were weakly correlated to genetic, but not to morphological differences. Our study is the first to reveal that both acoustic signaling and genetics clearly establish the species status for nocturnal primate populations. It also suggests that acoustic traits change at a more obvious and rapid pace than morphology in isolated populations, and may be used as an indication of conditions that may favor the evolution of subspecies.

Acoustics↗

Female dominance in captive gray mouse lemurs (Microcebus murinus).

Female dominance or female feeding priority seem to be characteristic for many lemur species, but are rare traits in other primates and mammals in general. The nocturnal lemur species, however, are underrepresented in the quantitative studies on social dominance. The aim of this study is to investigate the pattern of intersexual dominance relationships in the gray mouse lemur (Microcebus murinus), a species that is generally thought to possess a number of ancestral lemur traits. The context, distribution, and outcome of intersexual conflicts are analyzed in four captive groups of gray mouse lemurs. Intersexual conflicts occurred in the study groups in different behavioral contexts and were mostly spatial interactions (chasing/fleeing, approach/avoidance). The majority of conflicts were decided, and were in all but one case won by females. This is the first evidence suggesting unconditional female dominance in a cheirogaleid primate. The existence of female dominance in most families of the Lemuriformes suggests it is an ancient trait that evolved in their common ancestor.

Aggression↗

Behavioral study in the gray mouse lemur (Microcebus murinus) using compounds considered sweet by humans.

This study presents the results from two-bottle preference (TBP) tests performed on the gray mouse lemur (Microcebus murinus), a small Malagasy primate. We found that of 18 compounds considered sweet by humans, M. murinus preferred only six: D-tryptophan, dulcin, fructose, sucrose, SC45647, and xylitol. The animals neither preferred nor rejected acesulfame-K, alitame, aspartame, N-4-cyanophenyl-N'-cyanoguanidineacetate (CCGA), cyanosuosan, cyclamate, monellin, saccharin, suosan, super-aspartame, N-trifluoroacetyl-L-glutamyl-4-aminophenylcarbonitrile (TGC), and thaumatin. Together with previously recorded taste-nerve responses in M. murinus to acesulfame-K, alitame, aspartame, cyclamate, monellin, saccharin, and suosan [Hellekant et al., Chem Senses 18:307-320, 1993b], the current results suggest that these compounds either do not taste sweet to M. murinus or they have an aversive taste component. In this work we also relate these findings to phylogeny.

Animals↗

Comparative functional analysis of skull morphology of tree-gouging primates.

Many primates habitually feed on tree exudates such as gums and saps. Among these exudate feeders, Cebuella pygmaea, Callithrix spp., Phaner furcifer, and most likely Euoticus elegantulus elicit exudate flow by biting into trees with their anterior dentition. We define this behavior as gouging. Beyond the recent publication by Dumont ([1997] Am J Phys Anthropol 102:187-202), there have been few attempts to address whether any aspect of skull form in gouging primates relates to this specialized feeding behavior. However, many researchers have proposed that tree gouging results in larger bite force, larger internal skull loads, and larger jaw gapes in comparison to other chewing and biting behaviors. If true, then we might expect primate gougers to exhibit skull modifications that provide increased abilities to produce bite forces at the incisors, withstand loads in the skull, and/or generate large gapes for gouging. We develop 13 morphological predictions based on the expectation that gouging involves relatively large jaw forces and/or jaw gapes. We compare skull shapes for P. furcifer to five cheirogaleid taxa, E. elegantulus to six galagid species, and C. jacchus to two tamarin species, so as to assess whether gouging primates exhibit these predicted morphological shapes. Our results show little morphological evidence for increased force-production or load-resistance abilities in the skulls of these gouging primates. Conversely, these gougers tend to have skull shapes that are advantageous for creating large gapes. For example, all three gouging species have significantly lower condylar heights relative to the toothrow at a given mandibular length in comparison with closely related, nongouging taxa. Lowering the height of the condyle relative to the mandibular toothrow should reduce the stretching of the masseters and medial pterygoids during jaw opening, as well as position the mandibular incisors more anteriorly at wide jaw gapes. In other words, the lower incisors will follow a more vertical trajectory during both jaw opening and closing. We predict, based on these findings, that tree-gouging primates do not generate unusually large forces, but that they do use relatively large gapes during gouging. Of course, in vivo data on jaw forces and jaw gapes are required to reliably assess skull functions during gouging.

Anatomy, Comparative↗

Social organization of the golden brown mouse lemur (Microcebus ravelobensis).

Our study provides the first data on the social organization of the golden brown mouse lemur, a nocturnal primate discovered in northwestern Madagascar in 1994. The study was carried out in two 6-month field periods during the dry season, covering time before and during the mating season. The spatial and temporal distributions of the sexes in the population were investigated by mark/recapture and radiotelemetry. Focal observations and the determination of sleeping associations provided further insights into the sociality of this solitary forager. High intra- and intersexual home-range overlaps occurred throughout the study. In general, individuals of both sexes had spatial access to more than one conspecific of the same and the opposite sex. We found no indication for spatial monopolization of females by certain males. These results suggest a dispersed multimale/multifemale system with a promiscuous mating pattern. Individuals showed temporal stability in their home range locations and interacted regularly with conspecifics. Five sleeping groups were identified during the study period: one female group and four mixed-sex groups. Even though sleeping sites were changed frequently, sleeping-group compositions remained stable over time. Thermoregulatory constraints are the most likely explanation for sleeping-group composition with members of both sexes in this species. Mixed-sex sleeping groups can be described as the basic social unit within this dispersed multimale/multifemale society.

Animals↗

A shrew-sized origin for primates.

The origin of primates has had a long history of discussion and debate, with few ever considering the impact of the original body weight on subsequent primate adaptive radiations. Here, I attempt to reconstruct early primate evolution by considering the initial size of primates as well as the critical functional-adaptive events that had to occur prior to the early Eocene. Microcebus is often viewed as a living model, and thus 40-65 g might represent a practical ancestral weight for the origin of primates. I consider a smaller original body weight, likely 10-15 g in actual size, and I address the biological implications for shrew-sized primates by comparing the behavioral ecology of mouse lemurs, our smallest living primates, to another tiny-sized mammalian group, the shrews (Family Soricidae). Several behavioral and ecological characteristics are shared by shrews and mouse lemurs, and several mammalian trends are evident with decreased size. I suggest that a shrew-sized ancestral primate would have had high metabolic, reproductive, and predation rates, relatively low population densities, and a dispersed and solitary existence with a promiscuous mating system. Although small mammals like shrews provide insights concerning the ancestral size of primates, primate origins have always been tied to arboreality. I assess other potential arboreal models such as Ptilocercus and Caluromys. By combining all of this information, I try to sequence the events in a functional-adaptive series that had to occur before the early Eocene primate radiations. I suggest that all of these important adaptive events had to occur at a small body size below 50 g.

Adaptation, Physiological↗

Forelimb and hindlimb forces in walking and galloping primates.

One trait that distinguishes the walking gaits of most primates from those of most mammalian nonprimates is the distribution of weight between the forelimbs and hindlimbs. Nonprimate mammals generally experience higher vertical peak substrate reaction forces on the forelimb than on the hindlimb. Primates, in contrast, generally experience higher vertical peak substrate reaction forces on the hindlimb than on the forelimb. It is currently unclear whether this unusual pattern of force distribution characterizes other primate gaits as well. The available kinetic data for galloping primates are limited and present an ambiguous picture about peak-force distribution among the limbs. The present study investigates whether the pattern of forelimb-to-hindlimb force distribution seen during walking in primates is also displayed during galloping. Six species of primates were video-recorded during walking and galloping across a runway or horizontal pole instrumented with a force-plate. The results show that while the force differences between forelimb and hindlimb are not significantly different from zero during galloping, the pattern of force distribution is generally the same during walking and galloping for most primate species. These patterns and statistical results are similar to data collected during walking on the ground. The pattern of limb differentiation exhibited by primates during walking and galloping stands in contrast to the pattern seen in most nonprimate mammals, in which forelimb forces are significantly higher. The data reported here and by Demes et al. ([1994] J. Hum. Evol. 26:353-374) suggest that a relative reduction of forelimb vertical peak forces is part of an overall difference in locomotor mechanics between most primates and most nonprimate mammals during both walking and galloping.

Animals↗

Ontogeny of the nasopalatine duct in primates.

Ecological explanations have been put forward to account for the precocious or delayed development of patency in ducts leading to the vomeronasal organ (VNO) in certain mammals. Perinatal function may be related, in part, to the patency or fusion of the vomeronasal and nasopalatine (NPD) ducts. However, few studies have focused on NPD development in primates, which generally have a prolonged period of dependence during infancy. In this study we examined 24 prenatal primates and 13 neonatal primates, and a comparative sample of fetal mice and insectivores. In embryonic and early fetal Microcebus murinus, the NPD was completely fused, whereas in fetuses of later stages the duct was partially fused or completely patent. M. myoxinus of all stages demonstrated some degree of NPD fusion. In all other prenatal primates, the NPD was fused to some extent. Four prenatal insectivores (Tenrec ecaudatus) showed some degree of NPD fusion. In Mus musculus at 19 days gestation, the NPD was patent, although the anatomically separate VNO duct was fused. T. ecaudatus and most of the neonatal primates revealed complete NPD patency. An exception was Saguinus geoffroyi, which exhibited fusion of the NPD near the VNO opening. These observations may relate to differences in perinatal VNO function. The differences noted in our study suggest that M. murinus and M. myoxinus may differ in perinatal VNO functionality and perhaps in related behavior. Observations of neonatal primates suggest that NPD patency may be relatively common at birth and could serve other purposes in addition to being an access route for VNO stimuli.

Animals↗

Short and mid-wavelength cone distribution in a nocturnal Strepsirrhine primate (Microcebus murinus).

Strepsirrhines are of considerable interest for understanding the evolution of cone photoreceptors because they represent the most ancestral living primates. The retina of nocturnal Strepsirrhines is reported to contain a single population of medium/long wavelength (MW/LW) cones whereas short wavelength (SW) cones are totally absent. The area centralis of nocturnal Strepsirrhines also lacks the degree of central specialization seen in the fovea of diurnal primates. In this study of a nocturnal Strepsirrhine, the gray mouse lemur (Microcebus murinus), we used specific antibodies that recognize SW and MW/LW opsins to determine the presence of different cone subtypes and their distribution in relation to that of rods and ganglion cells. The results are compared to two diurnal Haplorhine species, a New World (Callithrix jacchus) and an Old World (Macaca fascicularis) monkey. In the mouse lemur, both antibodies to MW/LW cone opsin (COS-1 and CERN956) label the same population of cones. A small proportion of SW cones is only stained by the JH455 antiserum whereas the monoclonal OS-2 antibody shows negative staining. These two antibodies label the same SW cone population in other primates. The extracellular matrix of all cones is also labeled by the peanut agglutinin (PNA) lectin. In mouse lemur retinal wholemounts, peak cone density is localized at the area centralis and ranged from 7,500 to 8,000 cones/mm(2). SW cones represent less than 0.2 % of the total cone population and are mainly located in the nasal part of the retina. SW cones show an irregular distribution and densities never exceed 49 cones/mm(2). The distribution of neurons in the ganglion cell layer shows a distinct centroperipheral gradient with a peak of 28,000 cells/mm(2) at the area centralis. Rod distribution shows a centroperipheral gradient with the peak (850,000 rods/mm(2)) including and extending slightly dorsal to the area centralis. The theoretical spatial resolution of the mouse lemur (4.9 cycles/degree) is slightly lower to that of other nocturnal primates. The densities of rods, cones, and ganglion cell layer neurons represent a compromise between spatial resolution and sensitivity for both photopic and scotopic vision.

Animals↗

Seasonal variation in body mass and locomotor kinetics of the fat-tailed dwarf lemur (Cheirogaleus medius).

The fat-tailed dwarf lemur (Cheirogaleus medius) is unusual among primates in storing large amounts of fat subcutaneously prior to hibernating during the winter months. In doing so, it increases its body mass by more than 50%, with a substantial weight gain in the tail. This seasonal increase in mass provides a unique natural experiment to examine how changes in body mass affect substrate reaction forces during locomotion. As body mass increases, it is expected that the limbs of the fat-tailed dwarf lemur will be subjected to greater peak vertical substrate reaction forces during quadrupedal walking. However, whether or not these peak substrate reaction forces will increase proportionally across forelimbs and hindlimbs as body mass increases is unknown. Substrate reaction forces were collected on four adult C. medius walking quadrupedally on a 28-mm pole attached to a force platform. Peak vertical substrate reaction forces (Vpk) (N) were analyzed and compared for a cross-sectional sample of different body masses (180-300 g). Forelimb and hindlimb Vpk were positively correlated with body mass, with hindlimb Vpk always higher than forelimb Vpk. However, the rate at which Vpk increased relative to body mass was higher for the hindlimb than the forelimb. This disproportion in weight distribution between the forelimbs and hindlimbs as body mass increases appears to be linked to the accumulation of fat in the tail. It is likely that storing fat in the tail region may shift the center of mass more caudally, from a more cranial position when the tail is thinner. Such a caudal shift of the center of mass-either morphological or dynamic-is believed to have played an important role in the functional differentiation of the limbs and the evolution of locomotor modes of several tetrapod groups, including dinosaurs and primates.

Adipose Tissue↗

Regional atrophy in the brain of lissencephalic mouse lemur primates: measurement by automatic histogram-based segmentation of MR images.

Age-related regional cerebral atrophy was evaluated in a small lissencephalic primate, model of cerebral aging. Twelve mouse lemurs (Microcebus murinus), ages 1.9-10.9 years (maximum life span: 12 years), were studied. 3D inversion-recovery fast spin-echo MR images (isotropic resolution = 234 mum) were recorded at 4.7 T with a surface coil actively decoupled from the transmitting birdcage probe. The surface coil-related sensitivity gradient was corrected by normalization with images from an agar and NaCl phantom. An automatic statistical segmentation technique based on a classification-maximization algorithm was tested on digital phantoms that mimicked the brain and applied to the 3D brain images. Segmented 3D maps that displayed gray matter, white matter, and cerebro-spinal fluid (CSF) voxels were computed. The ventricles and peri-encephalic spaces were categorized into 14 regions, defined on brain atlases on the basis of cytoarchitectural and anatomical criteria. The volume of CSF voxels belonging to each of these regions was calculated as an index of regional atrophy. Dilation of the mammillary fossa was an early event in the aging process. CSF accumulation within the occipital, parieto-temporal, temporal, and frontal ventral peri-encephalic spaces was particularly marked in the oldest animals that also displayed ophthalmologic alterations.

Aging↗

Molecular cloning, sequencing, and brain expression of the presenilin 1 gene in Microcebus murinus.

The cDNA encoding the Microcebus murinus presenilin 1 protein (PSI) was cloned by RT-PCR from a brain cDNA library using various combinations of oligonucleotide primers designed on the basis of the human nucleotide sequence. Analysis of five clones isolated from two positive combinations revealed that the deduced open reading frame encodes two protein isoforms of 467 and 463 amino acid residues. The shorter isoform lacked the four residues VRSQ in the N-terminal region and like the 467 amino acid isoform presented 22 substitutions with its human homologue. The 12 bp nucleotide deletion evidenced in the cDNA encoding the shorter isoform is consistent with the use of an alternative 5' splice donor site identified at the end of the human exon 3. The immunohistochemistry performed with a specific polyclonal antiserum raised against a synthetic peptide located in the human large hydrophilic loop of PS1 revealed that the protein is widely distributed independently of age or of pathology in the microcebe brain. PS1 is found predominantly in neurons of the different cortical layers and hippocampus but also in subcortical structures. The PS1 labelling appeared as thin granulations scattered throughout the cytoplasm of numerous neurons and sometimes in neurites.

Aging↗

Cholinergic axonal dystrophy and mitochondrial pathology in prosimian primates.

Progressive cholinergic axonal dystrophy, cholinergic denervation, and generalized gliosis begin in the prosimian primate species Otolemur at 10% of maximum life span. In these same animals, extensive cerebral beta-amyloidosis follows relatively more abruptly at 50% of maximum life span. In contrast, even at maximum life span, the prosimian primate species Galago senegalensis Moholi, Microcebus murinus, and Eulemur fulvus collaris and insectivore species T. belangeri are either spared or much less affected. In this report, we further document this progressive cholinergic denervation in Otolemur which involves first projections of the pedunculopontine nucleus (PPN, CH5-6) and later projections of CH1-4 cholinergic nuclei, as well as other noncholinergic pathways. Affected cholinergic cell bodies and axons contain abnormal mitochondria with increased content of manganese superoxide dismutase (MnSOD). This syndrome correlates with moderate copper deficiency marked by diminished liver copper levels and cuproenzyme activities, carnitine deficiency possibly secondary to renal Fanconi syndrome, and evidence for stress inflammatory response activation. Mitochondrial pathology was observed in pancreatic islet cells, proximal renal tubule epithelial cells, and choroid plexus epithelial cells, and it involved central cholinergic neurons. In Otolemur garnetti, the degree of central cholinergic injury directly correlated to depression of liver copper stores. The Otolemur syndrome involves "sentinel" central cholinergic injury and selective mitochondrial pathology in cell classes defined by high mitochondrial content and/or metabolic activity and high content of nitric oxide synthetase and MnSOD. Environmental factors affecting copper and carnitine metabolism could interact with genetic defects or traits to produce abnormal and aggressive aging of Otolemur. Subclinical, cell-class specific mitochondrial dysfunction in these prosimian primates may be a model for human neurodegenerative diseases.

Aging↗

Identification of a unique apolipoprotein E allele in Microcebus murinus; ApoE brain distribution and co-localization with beta-amyloid and tau proteins.

This report is devoted to the characterization of the apolipoprotein E (ApoE) in Microcebus murinus. Only one allele homologous to the human ApoE4 allele was evidenced. The distribution of the corresponding ApoE protein in the brain was found in association with the pathological proteins characteristic of Alzheimer's disease (AD). Immunocytochemistry revealed brain deposits of ApoE in: (1) the cortical amyloid plaques; (2) the neurones of the various cortical lobes, the hippocampus and the brainstem; (3) the glial cells, astrocytes of the cortical parenchym and oligodendrocytes of the corpus callosum; and (4) the vessel walls. Most ApoE, beta-amyloid protein, abnormally phosphorylated Tau proteins and gliofilament acid proteins were seen in the same cortical areas. These findings for ApoE report the view that Microcebus murinus, in captivity, presents a pathological profile very similar to that observed in AD.

Alleles↗

beta-Amyloid protein precursor in Microcebus murinus: genotyping and brain localization.

Senile plaques characterized by beta-amyloid protein (A beta) deposits around dystrophic neurites and glial cells are more abundant in the cerebral cortical parenchyma of Alzheimer's disease (AD) patients than in the aged population. Four different mutations in the amyloid precursor protein (APP) gene have been directly involved in a few cases of familial AD with early onset (before 60 years). Previous studies have shown that Microcebus murinus, a nonhuman primate, also develops analogous deposits of A beta in the cortical parenchyma and blood vessel walls in the brain. Sequence analysis of exons 16 and 17 of the APP gene, encoding for A beta, revealed that even if nucleotide divergences occurred, the resulting peptide is completely homologous with the human A beta. The systematic comparison of the A beta nucleotide sequence in microcebus with or without amyloid deposits revealed that neither the presence of mutations involved in some cases of early onset familial AD nor the presence of a mutational founder effect can explain the amyloidosis observed in some old microcebus of our breeding. Localization of the APP was performed by immunocytochemistry in the brains of adult microcebus (1 to 11 years of age) using two antibodies raised against the C-terminus and N-terminus portions of APP. Microscopic examinations revealed that in the microcebus the APP distribution was similar to that observed in the human: (1) A beta and its precursor were simultaneously observed in amyloid plaques (AP) of the cortical parenchyma; (2) APP was localized in cell bodies and proximal dendrites of neurons, in astrocytes and oligodendrocytes, and in blood vessel and capillary walls; (3) labeling of APP in these structures was correlated with the presence of AP; and (4) labeling of APP increased with the age of the animal.

Amino Acid Sequence↗