Longicorn beetles (Coleoptera: Cerambycidae) differ considerably in the degree of their mushroom body development

A duality in the general structure of the mushroom body in longicorn beetles is confirmed. This duality is associated with the fact that they are formed by two solitary neuroblasts or two neuroblast clusters on each side of the brain and are manifested as a bipartite structure of both the calyx, which is the main sensory input, and the peduncular apparatus. Within the studied longicorn beetles, modifications in the general structure of mushroom bodies have been found; these modifications are caused by two oppositely directed morphogenetic processes, namely, the concentration of structures and their compartmentalization. The concentration leads to disappearance of the bipartite structure of the peduncular apparatus, whereas compartmentalization leads to a secondary subdivision of these structures into anatomically distinct subsections. This process is most pronounced in the peduncle and lobes. The mushroom bodies are best developed and differentiated in the members of the subfamily Lamiinae.     

Mushroom bodies in the brain of carrion beetles (Coleoptera, Silphidae)

Mushroom bodies are developed similarly in all the carrion beetles studied, and the transition from necrophagy to predation in some species does not affect considerably their level of development. The Kenyon cells form two small groups in each brain hemisphere and are subdivided into central and peripheral cells. Two closely located inputs of the Kenyon cell processes enter a single, pillow-shaped calyx region. The pedunculus comprises two shafts along a considerable part of its length, which fuse in the lobes. In the degree of development, the mushroom bodies of carrion beetles resemble those of basal lamellicorn and longhorned beetles.     

Not all dytiscidae have poorly developed mushroom bodies: The enigma of Cybister lateralimarginalis

The majority of diving beetles studied has completely differentiated but poorly developed mushroom bodies. The Kenyon cells are not numerous, the calyces are small, and the pedunculi and lobes have a simple structure. New Kenyon cells are produced by few solitary neuroblasts. Cybister lateralimarginalis makes an amazing exception. Its mushroom bodies are strongly developed and comprise numerous Kenyon cells, large calyces, and a peduncular apparatus of a complicated structure. The Kenyon cells are produced in polyneuroblast proliferative centers. The grounds of such strong development of the mushroom body in Cybister remain unknown.     

Histological structure of tripartite mushroom bodies in ground beetles (Insecta, Coleoptera: Carabidae)

Contrary to members of the suborder Polyphaga, ground beetles have been found to possess tripartite mushroom bodies, which are poorly developed in members of basal taxa and maximally elaborated in evolutionarily advanced groups. Nevertheless, they do not reach the developmental stage, which has been previously found in particular families of beetles. It has been pointed out that a new formation of the Kenyon cells occurs during at least the first months of adult life, and inactive neuroblasts are found even in one-year-old beetles. It has been suggested that there is a relation between the Kenyon cell number and development of the centers of Kenyon cell new-formation.     

Some cetoniinae (Coleoptera,Scarabaeidae) have structurally different medial and lateral calyces of mushroom bodies

Structural differences between the medial and lateral calyces of mushroom bodies in insects are described for the first time. In two cetoniine scarab beetles, Cetonia aurata and Oxythyrea funesta, the lateral calyces are subdivided into two portions showing a different neuropil structure. This feature is not reflected in the structure of the pedunculus and lobes, as well as in the relative neuropil volume occupied by transformed lateral calyx as compared with unmodified lateral calyx of related scarab beetles. The lateral calyx modification is considered to be related to changes in dendritic arborizations of central Kenyon cells. The subdivision of lateral calyx occurs only in adults and was not observed in larvae.     

A subpopulation of mushroom body intrinsic neurons is generated by protocerebral neuroblasts in the tobacco hornworm moth, Manduca sexta (Sphingidae, Lepidoptera)

Subpopulations of Kenyon cells, the intrinsic neurons of the insect mushroom bodies, are typically sequentially generated by dedicated neuroblasts that begin proliferating during embryogenesis. When present, Class III Kenyon cells are thought to be the first born population of neurons by virtue of the location of their cell somata, farthest from the position of the mushroom body neuroblasts. In the adult tobacco hornworm moth Manduca sexta, the axons of Class III Kenyon cells form a separate Y tract and dorsal and ventral lobelet; surprisingly, these distinctive structures are absent from the larval Manduca mushroom bodies. BrdU labeling and immunohistochemical staining reveal that Class III Kenyon cells are in fact born in the mid-larval through adult stages. The peripheral position of their cell bodies is due to their genesis from two previously undescribed protocerebral neuroblasts distinct from the mushroom body neuroblasts that generate the other Kenyon cell types. These findings challenge the notion that all Kenyon cells are produced solely by the mushroom body neuroblasts, and may explain why Class III Kenyon cells are found sporadically across the insects, suggesting that when present, they may arise through de novo recruitment of neuroblasts outside of the mushroom bodies. In addition, lifelong neurogenesis by both the Class III neuroblasts and the mushroom body neuroblasts was observed, raising the possibility that adult neurogenesis may play a role in mushroom body function in Manduca.     

Developmental organization of the mushroom bodies of Thermobia domestica (Zygentoma, Lepismatidae): insights into mushroom body evolution from a basal insect

The mushroom bodies of the insect brain are sensory integration centers best studied for their role in learning and memory. Studies of mushroom body structure and development in neopteran insects have revealed conserved morphogenetic mechanisms. The sequential production of morphologically distinct intrinsic neuron (Kenyon cell) subpopulations by mushroom body neuroblasts and the integration of newborn neurons via a discrete ingrowth tract results in an age-based organization of modular subunits in the primary output neuropil of the mushroom bodies, the lobes. To determine whether these may represent ancestral characteristics, the present account assesses mushroom body organization and development in the basal wingless insect Thermobia domestica. In this insect, a single calyx supplied by the progeny of two neuroblast clusters, and three perpendicularly oriented lobes are readily identifiable. The lobes are subdivided into 15 globular subdivisions (Trauben). Lifelong neurogenesis is observed, with axons of newborn Kenyon cells entering the lobes via an ingrowth core. The Trauben do not appear progressively during development, indicating that they do not represent the ramifications of sequentially produced subpopulations of Kenyon cells. Instead, a single Kenyon cell population produces highly branched axons that supply all lobe subdivisions. This suggests that although the ground plan for neopteran mushroom bodies existed in early insects, the organization of modular subunits composed of separate Kenyon cell subpopulations is a later innovation. Similarities between the calyx of Thermobia and the highly derived fruit fly Drosophila melanogaster also suggest a correlation between calyx morphology and Kenyon cell number.     

<Emphasis Type="Italic">Amphicoma</Emphasis> (Coleoptera,Glaphyridae): A generalist feeder with poorly developed mushroom bodies

Species of the genus Amphicoma are phytophagous generalists but have poorly developed mushroom bodies with a minute calyx and single (undivided) pedunculus and lobes of concentric structure. Thus, in this respect Amphicoma are closer to the dung scarab beetles regarded as specialist feeders (Farris and Roberts, 2005).     

Structure of the mushroom bodies in Scarabaeoidea (Insecta: Coleoptera): 1. Basal families and coprophagous scarabaeidae

Mushroom bodies are in general similarly developed in most taxons studied. The calyx region appears as a single structure, and its dual nature is not yet realized. An anterio-posterior asymmetry of the calyx region with Kenyon cell processes running mostly behind the glomerular neuropil of the calyx is characteristic of all the species studied. In this respect, the calyx region of basal Scarabaeoidea resembles greatly the calyx of many dipterans. Lobe compartmentalization occurs at the initial stage. The passalid beetles represent an exception, as their mushroom bodies are much more developed than in related families. This may be connected with the complicated social behavior of Passalidae.     

Separate distribution of deutocerebral projection neurons in the mushroom bodies of the cricket brain

Deutocerebral projection neurones in the brain of the cricket (Gryllus bimaculatus) have been investigated by experimental dextran staining, viewed by light and electron microscopy. These neurones of two separate somata clusters innervate two separate primary glomerular neuropils of the deutocerebral segment, either the antennal lobe receiving only antennal nerve sensory input, or the glomerular lobe, receiving input from sensory neurones of lower segmental origin, including chemosensory fibres from mouth parts. Projection neurones of the antennal lobe only invade the anterior calyx of the mushroom body neuropil via the inner antenno glomerular tract, while glomerular relay neurones of the glomerular lobe innervate only the posterior calyx via the tritocerebral tract. All types of projection neurones give rise to presynaptic boutons. forming the central core of microglomeruli with patterned distribution. These projection neurons are cholinergic. The results are discussed in view of maintained segregated modal information, first processed in the separated primary deutocerebral neuropiles and further on in the second order input neuropils of the mushroom bodies. The large posterior calyces are proposed as a compartment for gustatory information.     

Behavioral functions of the insect mushroom bodies

New methods of intervention in Drosophila and other insect species reveal that the mushroom bodies are involved in a diverse set of behavioral functions. The intrinsic Kenyon cells (those neurons with projections within the mushroom bodies) house part of the short-term memory trace for odors and are required for courtship conditioning memory. A pair of extrinsic mushroom body neurons (neurons with projections both inside and outside the mushroom bodies) provides a neuropeptide important for 1-hour olfactory memory. In addition, the mushroom bodies are necessary for context generalization in visual learning and for regulating the transition from walking to rest.     

Brain central regions in courtship sound production in Drosophila melanogaster

There are debates about the function of the two main central brain structures of insects--mushroom bodies and the central complex--in the control of motor co-ordination and triggering of different behaviour programs including sound production. To throw additional light onto this problem we analysed the parameters of the love song produced by 5-day old males courting for 5 minutes a fertilised CS female at 25 degrees C, in two wild-type strains of Drosophila melanogaster (Berlin and CS), hydroxyurea (HU)-treated flies (chemical ablation of the mushroom bodies) two mushroom body mutants (mbm1 and mud1), two central complex mutants (ccbKS127 and cexKS181) and a mutant cxbN71 with defects both in the mushroom bodies and in the central complex. It was found that the love song of HU-treated flies devoid of the mushroom bodies is very similar to that of wild-type flies. In mbm1 and mud1 the main parameters of the song (interpulse interval, IPI, and train duration) are slightly shifted from those of wild type but the sharpness of tuning of the pulse oscillator is the same. The flies of all these strains are equal to wild-type strains in mating success (% of copulations with virgins in 10-min test). On the contrary, the songs of the central complex mutants differ from those of wild-type flies. First of all, the sharpness of tuning of the pulse oscillator is destroyed,--the IPIs become highly variable. The pulses often are much longer and polycyclic as in well known cacophony mutant. The mean duration of pulse trains is much shorter. The males of the mutant cexKS181 usually court violently, but in most cases abnormal sounds are produced. Both cexKS181 and ccbKS127 males are much less successful in matings in comparison to wild-type flies. One can conclude that the central complex plays probably a very important role in the control of singing, whereas the mushroom bodies are practically not involved in this function.     

草原珍稀食用菌蒙古口蘑菌种的分子鉴定

目的:蒙古口蘑是一种著名的野生食用菌.为了辨别蒙古口蘑与其相似的草原蘑菇种及分离菌株的真伪.方法:采用ITS序列分析方法对从内蒙古地区采集的子实体和纯培养菌株进行分子鉴定.结果:在20个供试材料中,从呼伦贝尔草原采集的4个子实体(来源于不同地区)中有2个属于蒙古口蘑,从子实体分离得到的纯培养物中有2个为蒙古口蘑菌株;在呼伦贝尔地区购买的蘑菇干品和冷冻品都属于蒙古口蘑;从锡林郭勒草原的不同地区采集分离得到的10株纯培养物中有6株为蒙古口蘑菌株.结论:在呼伦贝尔地区和锡盟地区都有蒙古口蘑存在,而且从不同地区采集的蒙古口蘑亲缘关系比较近.     

Global and local modulatory supply to the mushroom bodies of the moth Spodoptera littoralis

The moth Spodoptera littoralis, is a major pest of agriculture whose olfactory system is tuned to odorants emitted by host plants and conspecifics. As in other insects, the paired mushroom bodies are thought to play pivotal roles in behaviors that are elicited by contextual and multisensory signals, amongst which those of specific odors dominate. Compared with species that have elaborate behavioral repertoires, such as the honey bee Apis mellifera or the cockroach Periplaneta americana, the mushroom bodies of S. littoralis were originally viewed as having a simple cellular organization. This has been since challenged by observations of putative transmitters and neuromodulators. As revealed by immunocytology, the spodopteran mushroom bodies, like those of other taxa, are subdivided longitudinally into discrete neuropil domains. Such divisions are further supported by the present study, which also demonstrates discrete affinities to different mushroom body neuropils by antibodies raised against two putative transmitters, glutamate and gamma-aminobutyric acid, and against three putative neuromodulatory substances: serotonin, A-type allatostatin, and tachykinin-related peptides. The results suggest that in addition to longitudinal divisions of the lobes, circuits in the calyces and lobes are likely to be independently modulated.     

Evolution of insect mushroom bodies: old clues,new insights

The mushroom bodies are a morphologically diverse sensory integration and learning and memory center in the brains of various invertebrate species, of which those of insects are the best described. Insect mushroom bodies are composed of numerous tiny intrinsic neurons (Kenyon cells) that form calyces with their dendrites and a pedunculus and lobes with their axons. The identities of conserved Kenyon cell subpopulations and the correlations between morphological and functional specializations of the mushroom bodies are just beginning to be elucidated, providing insight into mechanisms of mushroom body evolution. Comparisons of mushroom body organization in different insect lineages reveal trends in the evolution of subcompartments correlated with the elaboration, reduction, acquisition or loss of Kenyon cell subpopulations. Furthermore, these changes often appear correlated with variation in type and strength of afferent input and in behavioral ecology. These and other features of mushroom body organization suggest a striking convergence with mammalian cortex, with Kenyon cell subpopulations displaying evolutionary modularity in a manner reminiscent of cortical areas.     

Are mushroom bodies cerebellum-like structures?

The mushroom bodies are distinctive neuropils in the protocerebral brain segments of many protostomes. A defining feature of mushroom bodies is their intrinsic neurons, masses of cytoplasm-poor globuli cells that form a system of lobes with their densely-packed, parallel-projecting axon-like processes. In insects, the role of the mushroom bodies in olfactory processing and associative learning and memory has been studied in depth, but several lines of evidence suggest that the function of these higher brain centers cannot be restricted to these roles. The present account considers whether insight into an underlying function of mushroom bodies may be provided by cerebellum-like structures in vertebrates, which are similarly defined by the presence of masses of tiny granule cells that emit thin parallel fibers forming a dense molecular layer. In vertebrates, the shared neuroarchitecture of cerebellum-like structures has been suggested to underlie a common functional role as adaptive filters for the removal of predictable sensory elements, such as those arising from reafference, from the total sensory input. Cerebellum-like structures include the vertebrate cerebellum, the electrosensory lateral line lobe, dorsal and medial octavolateral nuclei of fish, and the dorsal cochlear nucleus of mammals. The many architectural and physiological features that the insect mushroom bodies share with cerebellum-like structures suggest that it might be fruitful to consider mushroom body function in light of a possible role as adaptive sensory filters. The present account thus presents a detailed comparison of the insect mushroom bodies with vertebrate cerebellum-like structures.     

Experience-dependent plasticity in the mushroom bodies of the solitary bee Osmia lignaria (Megachilidae)

All members of the solitary bee species Osmia lignaria (the orchard bee) forage upon emergence from their natal nest cell. Conversely, in the honey bee, days-to-weeks of socially regulated behavioral development precede the onset of foraging. The social honey bee's behavioral transition to foraging is accompanied by neuroanatomical changes in the mushroom bodies, a region of the insect brain implicated in learning. If these changes were general adaptations to foraging, they should also occur in the solitary orchard bee. Using unbiased stereological methods, we estimated the volume of the major compartments of the mushroom bodies, the neuropil and Kenyon cell body region, in adult orchard bees. We compared the mushroom bodies of recently emerged bees with mature bees that had extensive foraging experience. To separate effects of general maturation from field foraging, some orchard bees were confined to a cage indoors. The mushroom body neuropil of experienced field foragers was significantly greater than that of both recently emerged and mature caged orchard bees, suggesting that, like the honey bee, this increase is driven by outdoor foraging experience. Unlike the honey bee, where increases in the ratio of neuropil to Kenyon cell region occur in the worker after emerging from the hive cell, the orchard bee emerged from the natal nest cell with a ratio that did not change with maturation and was comparable to honey-bee foragers. These results suggest that a common developmental endpoint may be reached via different development paths in social and solitary species of foraging bees.     

Higher order visual input to the mushroom bodies in the bee, Bombus impatiens

To produce appropriate behaviors based on biologically relevant associations, sensory pathways conveying different modalities are integrated by higher-order central brain structures, such as insect mushroom bodies. To address this function of sensory integration, we characterized the structure and response of optic lobe (OL) neurons projecting to the calyces of the mushroom bodies in bees. Bees are well known for their visual learning and memory capabilities and their brains possess major direct visual input from the optic lobes to the mushroom bodies. To functionally characterize these visual inputs to the mushroom bodies, we recorded intracellularly from neurons in bumblebees (Apidae: Bombus impatiens) and a single neuron in a honeybee (Apidae: Apis mellifera) while presenting color and motion stimuli. All of the mushroom body input neurons were color sensitive while a subset was motion sensitive. Additionally, most of the mushroom body input neurons would respond to the first, but not to subsequent, presentations of repeated stimuli. In general, the medulla or lobula neurons projecting to the calyx signaled specific chromatic, temporal, and motion features of the visual world to the mushroom bodies, which included sensory information required for the biologically relevant associations bees form during foraging tasks.     

Influence of spatio-temporal resource availability on mushroom mite diversity

Although biodiversity in nature is of fundamental importance because it improves the sustainability of ecosystems, communities of microscopic organisms are generally excluded from conservation targets for biodiversity. Here, I hypothesize that mushroom mite species richness is correlated with both spatial (i.e., mushroom size) and temporal (i.e., longevity of fruiting bodies) resource availability. I collected fruiting bodies in an old-growth forest over 4 years to collect mites and insects inhabiting the mushrooms. Mites were collected from 47 % of the fruiting bodies and approximately 60 % of the mite species were collected only once. Mite species richness was significantly correlated with the availability of long-lasting fruiting bodies. For example, bracket fungi contained more mite species than ephemeral fruiting bodies. Insect presence was also correlated with mushroom mite richness, probably as phoretic hosts and food resources for predacious mites. On the other hand, mushroom size seemed to be less important; small fruiting bodies sometimes harbored several mite species. Although mite species richness was correlated with mushroom species richness, mushroom specificity by mites was not clear except for a preference for long-lasting fruiting bodies. Therefore, I suggest that a constant supply of coarse woody debris is crucial for maintaining preferred resources for mushroom mites (e.g., bracket fungi) and their associated insects (mycophilous and possibly saproxylic insects).     

The mushroom bodies of the lower nematocera: A link between those of the higher diptera and other mecopteroids

Nematoceran Diptera are nonuniform in the structure of their mushroom bodies. Members of the more basal families (Ptychopteridae, Pediciidae, and Tipulidae) have bipartite mushroom bodies, characteristic of members of the other mecopteroid complex orders. In members of Bibionomorpha (Bibionidae and Anisopodidae), tripartite mushroom bodies have been found characteristic of Brachycera Orthorrhapha.