The sense of smell,its signalling pathways,and the dichotomy of cilia and microvilli in olfactory sensory cells

Smell is often regarded as an ancillary perception in primates, who seem so dominated by their sense of vision. In this paper, we will portray some aspects of the significance of olfaction to human life and speculate on what evolutionary factors contribute to keeping it alive. We then outline the functional architecture of olfactory sensory neurons and their signal transduction pathways, which are the primary detectors that render olfactory perception possible. Throughout the phylogenetic tree, olfactory neurons, at their apical tip, are either decorated with cilia or with microvilli. The significance of this dichotomy is unknown. It is generally assumed that mammalian olfactory neurons are of the ciliary type only. The existence of so-called olfactory microvillar cells in mammals, however, is well documented, but their nature remains unclear and their function orphaned. This paper discusses the possibility, that in the main olfactory epithelium of mammals ciliated and microvillar sensory cells exist concurrently. We review evidence related to this hypothesis and ask, what function olfactory microvillar cells might have and what signalling mechanisms they use.     

Factors that regulate embryonic gustatory development

Smell is often regarded as an ancillary perception in primates, who seem so dominated by their sense of vision. In this paper, we will portray some aspects of the significance of olfaction to human life and speculate on what evolutionary factors contribute to keeping it alive. We then outline the functional architecture of olfactory sensory neurons and their signal transduction pathways, which are the primary detectors that render olfactory perception possible. Throughout the phylogenetic tree, olfactory neurons, at their apical tip, are either decorated with cilia or with microvilli. The significance of this dichotomy is unknown. It is generally assumed that mammalian olfactory neurons are of the ciliary type only. The existance of so-called olfactory microvillar cells in mammals, however, is well documented, but their nature remains unclear and their function orphaned. This paper discusses the possibility, that in the main olfactory epithelium of mammals ciliated and microvillar sensory cells exist concurrently. We review evidence related to this hypothesis and ask, what function olfactory microvillar cells might have and what signalling mechanisms they use.

     

Magnetohydrodynamic waves

Smell is a physical process used by us all, but fully understood by none. A physicist seeks to understand the mysterious phenomena of nature and so it is natural to apply physics to this problem. There is a reluctance in the field of olfactory science to explore the physical processes of fundamental interactions in odorant recognition because the system is complex. However, nature is often compliant to our simple models, and part of the art is to reduce the problem to a soluble one, and part of the beauty of science is often that the simple model with the simple answers give the right ones. This article reviews the applicability of some physical models to olfaction and looks in particular at one theory within the realm of olfactory science. This theory was first endorsed by Turin in 1996, that smell signalling may be based on a form of Inelastic Electron Tunnelling (IETS). This review looks at the mystery of olfaction and the physical principles that may unveil such mysterious phenomena.     

基于气体浓度定量的嗅觉刺激器优化设计

现有的基于磁共振测量的嗅觉刺激器，通过调节嗅剂液体浓度的方法可以实现不同浓度的嗅觉刺激，但随着实验进行，受到嗅剂挥发以及实验环境（温度、湿度、气流量）变化的影响，很难确保输送至鼻腔的嗅剂气体浓度的稳定性，进而影响实验结果的准确性．本研究对本实验室前期开发的嗅觉刺激装置进行改进，实现了气体浓度精确定量．改进后的嗅觉刺激器主要分为三个部分：控制系统、反馈系统和气路系统．控制系统主要实现气路系统的送气控制和嗅剂气体浓度调节；反馈系统则负责对气体浓度进行测量；气路系统则在原有基础上添加活性炭装置，降低无关因素干扰．装置改进之后，不同气路切换时间为75.2 ms，比原装置减少了1 s，有效提高刺激精度．实验结果显示，气体浓度调节前，300 s内乙醇、吡啶、乙酸戊酯嗅剂气体浓度分别下降6.7%、71.4%、79.2%，嗅剂气体浓度短时间内发生较大改变．加入气体浓度调节功能后，当气体浓度下降至目标浓度的90%时，可通过调节气泵电压改变嗅剂气流与空气气流比例，从而调节嗅剂气体浓度至目标值，其中吡啶、乙酸戊酯用时13 s．     

Brain architecture in the terrestrial hermit crab Coenobita clypeatus (Anomura, Coenobitidae), a crustacean with a good aerial sense of smell

Background

During the evolutionary radiation of Crustacea, several lineages in this taxon convergently succeeded in meeting the physiological challenges connected to establishing a fully terrestrial life style. These physiological adaptations include the need for sensory organs of terrestrial species to function in air rather than in water. Previous behavioral and neuroethological studies have provided solid evidence that the land hermit crabs (Coenobitidae, Anomura) are a group of crustaceans that have evolved a good sense of aerial olfaction during the conquest of land. We wanted to study the central olfactory processing areas in the brains of these organisms and to that end analyzed the brain of Coenobita clypeatus (Herbst, 1791; Anomura, Coenobitidae), a fully terrestrial tropical hermit crab, by immunohistochemistry against synaptic proteins, serotonin, FMRFamide-related peptides, and glutamine synthetase.

Results

The primary olfactory centers in this species dominate the brain and are composed of many elongate olfactory glomeruli. The secondary olfactory centers that receive an input from olfactory projection neurons are almost equally large as the olfactory lobes and are organized into parallel neuropil lamellae. The architecture of the optic neuropils and those areas associated with antenna two suggest that C. clypeatus has visual and mechanosensory skills that are comparable to those of marine Crustacea.

Conclusion

In parallel to previous behavioral findings of a good sense of aerial olfaction in C. clypeatus, our results indicate that in fact their central olfactory pathway is most prominent, indicating that olfaction is a major sensory modality that these brains process. Interestingly, the secondary olfactory neuropils of insects, the mushroom bodies, also display a layered structure (vertical and medial lobes), superficially similar to the lamellae in the secondary olfactory centers of C. clypeatus. More detailed analyses with additional markers will be necessary to explore the question if these similarities have evolved convergently with the establishment of superb aerial olfactory abilities or if this design goes back to a shared principle in the common ancestor of Crustacea and Hexapoda.     

脑功能磁共振成像在人类嗅觉研究中的应用

在人类的5种主要感觉中,嗅觉是最广泛、古老、直接和内在的感觉.这些特性使人们对人类嗅觉的研究异常艰难,以致于直到今天人们对嗅觉的功能仍不清楚,而对大脑的功能机制所知更少.与其他基于物理原理的方法一样,磁共振成像技术的广泛应用极大地推动了整个生命科学的发展.脑功能磁共振成像的优势(高分辨率、高对比度、无损性和无放射性等)为人们研究嗅觉高级中枢以及与嗅觉相关行为的脑机制等提供了强有力的技术手段.文章在简单介绍嗅觉知识的基础上,着重讨论了近十年来,脑功能磁共振成像技术在人类嗅觉研究中所取得的成果.     

Tiling solutions for optimal biological sensing

Biological systems, from cells to organisms, must respond to the ever-changing environment in order to survive and function. This is not a simple task given the often random nature of the signals they receive, as well as the intrinsically stochastic, many-body and often self-organized nature of the processes that control their sensing and response and limited resources. Despite a wide range of scales and functions that can be observed in the living world, some common principles that govern the behavior of biological systems emerge. Here I review two examples of very different biological problems: information transmission in gene regulatory networks and diversity of adaptive immune receptor repertoires that protect us from pathogens. I discuss the trade-offs that physical laws impose on these systems and show that the optimal designs of both immune repertoires and gene regulatory networks display similar discrete tiling structures. These solutions rely on locally non-overlapping placements of the responding elements (genes and receptors) that, overall, cover space nearly uniformly.     

Mechanism of smell: Electrochemistry,receptors and cell signaling

This report presents in four main parts a novel approach based on electrochemistry, receptors and cell signaling. In Part A, there is limited correlation between dipole moments (DMs), associated electrostatic fields (EFs), and odor. For Part B, binding of the odorant to the receptor results in interaction of the ligand EF with those of the protein receptor, resulting in alteration. Part C addresses passage of the message by the altered EF to the olfactory neurons. Part D represents the final step in which the electrical signal is converted to perceived odor in the cerebral olfactory cortex.     

Analyzing neuronal networks using discrete-time dynamics

We develop mathematical techniques for analyzing detailed Hodgkin-Huxley like models for excitatory-inhibitory neuronal networks. Our strategy for studying a given network is to first reduce it to a discrete-time dynamical system. The discrete model is considerably easier to analyze, both mathematically and computationally, and parameters in the discrete model correspond directly to parameters in the original system of differential equations. While these networks arise in many important applications, a primary focus of this paper is to better understand mechanisms that underlie temporally dynamic responses in early processing of olfactory sensory information. The models presented here exhibit several properties that have been described for olfactory codes in an insect’s Antennal Lobe. These include transient patterns of synchronization and decorrelation of sensory inputs. By reducing the model to a discrete system, we are able to systematically study how properties of the dynamics, including the complex structure of the transients and attractors, depend on factors related to connectivity and the intrinsic and synaptic properties of cells within the network.     

Gap junctions in olfactory neurons modulate olfactory sensitivity

Background  

One of the fundamental questions in olfaction is whether olfactory receptor neurons (ORNs) behave as independent entities within the olfactory epithelium. On the basis that mature ORNs express multiple connexins, I postulated that gap junctional communication modulates olfactory responses in the periphery and that disruption of gap junctions in ORNs reduces olfactory sensitivity. The data collected from characterizing connexin 43 (Cx43) dominant negative transgenic mice OlfDNCX, and from calcium imaging of wild type mice (WT) support my hypothesis.     

Investigation of organic films by atomic force microscopy: Structural,nanotribological and electrical properties

Atomic force microscopy (AFM) has found its applications in a wide range of research fields. In this review, we show by examples that atomic force microscopy is a powerful technique to investigate structural, mechanical and electrical properties of organic films. We start with an introduction of AFM instrumentation highlighting AFM developments that are of direct relevance to organic films. Next, we review AFM studies on organic films according to their preparation methods: self-assembly, the Langmuir–Blodgett technique, solution preparation, and thermal evaporation. In the discussion on self-assembled monolayers, we focus on aspects such as structural evolution, load-induced molecular tilting, annealing, and incorporation of conjugated groups. For solution prepared organic films, we stress annealing-induced structural evolution as well as the effects of phase separation/segregation. We also briefly summarize the progress of AFM investigation on Langmuir–Blodgett films and thermally evaporated organic films. We conclude the review by providing some thoughts for future exploration. In particular, atomic force microscopy combined with ultra-flat coplanar nano-electrodes provides a promising platform to isolate single or a small number of molecular features (e.g. vacancies, defects, grain boundaries) in organic films as well as to identify the role of these features at the nanometer scale.     

一种用于fMRI的嗅觉刺激装置优化与验证

为搭建可用于磁共振环境下的自动控制嗅觉刺激器,本文根据刺激装置搭建的通用要求和实验室已有装置的性能提出了改进需求和系统整体设计方案．刺激装置分为控制系统与气体输送系统两部分．控制系统的软件部分基于LabVIEW平台编程,采用了虚拟仪器方案,提供可输入刺激序列的人机界面,并根据不同的刺激需求来控制电磁阀动作,以切换不同气路．气体输送系统由4条可变支路和1条恒流支路组成,其中的3条可变气路由洁净空气分别通过装有不同气味液体的洗气瓶来产生3种刺激气味．系统搭建完成后,使用霍尼韦尔AWM43600空气流量传感器测量了系统气体流量波动率为0.3%,同时测得不同刺激气路切换时的切换响应时间为1.07 s．最后使用该刺激装置对8名被试进行嗅觉刺激的同时进行功能磁共振成像（fMRI）实验,实验采用了乙醇、吡啶和乙酸异戊酯3种刺激气味,fMRI图像结果显示被试的嗅觉受到刺激后,丘脑、杏仁核、梨状皮质、眶额皮层等嗅觉相关脑区激活．以上实验表明,本文搭建的指标可量化的刺激器更能满足嗅觉fMRI实验的要求．     

Olfactory and solitary chemosensory cells: two different chemosensory systems in the nasal cavity of the American alligator, <Emphasis Type="Italic">Alligator mississippiensis</Emphasis>

Background  

The nasal cavity of all vertebrates houses multiple chemosensors, either innervated by the Ist (olfactory) or the Vth (trigeminal) cranial nerve. Various types of receptor cells are present, either segregated in different compartments (e.g. in rodents) or mingled in one epithelium (e.g. fish). In addition, solitary chemosensory cells have been reported for several species. Alligators which seek their prey both above and under water have only one nasal compartment. Information about their olfactory epithelium is limited. Since alligators seem to detect both volatile and water-soluble odour cues, I tested whether different sensory cell types are present in the olfactory epithelium.     

High-pressure properties of group IV clathrates

We review the investigations on clathrate materials having C, Si, Ge or Sn framework atoms in which the pressure parameter plays an important role. In this article, we studied the synthesis of clathrates, superconducting properties and those studies relative to the structural cohesivity and phase diagrams. We have tried to extensively review these subjects. Key references on other important properties of the group IV clathrates are provided in the introductory sections. As a main result of this review, we note that pressure appears as a key parameter for the elaboration of present and future clathrates in particular those exhibiting superconducting properties. We show how high-pressure research has also played an important role in the understanding of the parameters governing clathrate superconductivity. In contrast, the study of the structural evolution of group IV clathrates (bulk modulus and stability) gives abundant clues for tailoring new materials with improved mechanical properties. Finally, practically all investigations converge to point out that today hypothetical carbon clathrates are candidates for extraordinary superconducting and mechanical properties.     

Statistical Mechanics of Horizontal Gene Transfer in?Evolutionary Ecology

The biological world, especially its majority microbial component, is strongly interacting and may be dominated by collective effects. In this review, we provide a brief introduction for statistical physicists of the way in which living cells communicate genetically through transferred genes, as well as the ways in which they can reorganize their genomes in response to environmental pressure. We discuss how genome evolution can be thought of as related to the physical phenomenon of annealing, and describe the sense in which genomes can be said to exhibit an analogue of information entropy. As a direct application of these ideas, we analyze the variation with ocean depth of transposons in marine microbial genomes, predicting trends that are consistent with recent observations using metagenomic surveys.     

TRPM5-expressing microvillous cells in the main olfactory epithelium

Background  

The main olfactory epithelium (MOE) in the nasal cavity detects a variety of air borne molecules that provide information regarding the presence of food, predators and other relevant social and environmental factors. Within the epithelium are ciliated sensory neurons, supporting cells, basal cells and microvillous cells, each of which is distinct in morphology and function. Arguably, the least understood, are the microvillous cells, a population of cells that are small in number and whose function is not known. We previously found that in a mouse strain in which the TRPM5 promoter drives expression of the green fluorescent protein (GFP), a population of ciliated olfactory sensory neurons (OSNs), as well as a population of cells displaying microvilli-like structures is labeled. Here we examined the morphology and immunocytochemical properties of these microvillous-like cells using immunocytochemical methods.     

Effects of aversive odour presentation on inhibitory control in the Stroop colour-word interference task

Background  

Due to the unique neural projections of the olfactory system, odours have the ability to directly influence affective processes. Furthermore, it has been shown that emotional states can influence various non-emotional cognitive tasks, such as memory and planning. However, the link between emotional and cognitive processes is still not fully understood. The present study used the olfactory pathway to induce a negative emotional state in humans to investigate its effect on inhibitory control performance in a standard, single-trial manual Stroop colour-word interference task. An unpleasant (H₂S) and an emotionally neutral (Eugenol) odorant were presented in two separate experimental runs, both in blocks alternating with ambient air, to 25 healthy volunteers, while they performed the cognitive task.     

Role of quantum fluctuations in a system with strong fields: Spectral properties and thermalization

In a previous work [arXiv:1009.4363], we have studied the evolution of a scalar field with a quartic coupling, driven by a classical source that initializes it to a non-perturbatively large value. At leading order in the coupling, the evolution of this system is given by classical solutions of the field equation of motion. However, this system is subject to a parametric resonance that leads to secular divergences in higher-order corrections to physical observables. We have proposed a scheme that resums all the leading secular terms: this resummation leads to finite results at all times, and we have observed also that it makes the pressure tensor of the system relax to its equilibrium value.In the present paper, we continue the study of this system by looking at finer details of its dynamics. We first compute its spectral function at various stages of the evolution, and we observe that after a fairly short transient time there are well-defined massive quasi-particles. We then consider the time evolution of the momentum distribution of these quasi-particles, and we show that after a stage dominated by the parametric resonance, this distribution slowly evolves to an equilibrium distribution. Interestingly, this distribution develops a transient chemical potential, signaling the fact that number changing processes are much slower than the elastic ones.