Functional and structural mapping of human cerebral cortex: solutions are in the surfaces
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Radiation exposures to individuals in space can greatly exceed natural radiation exposure on Earth and possibly normal occupational radiation exposures as well. Consequently, procedures limiting exposures would be necessary. Limitations were proposed by the Radiobiological Advisory Panel of the National Academy of Sciences/National Research Council in 1970. This panel recommended short-term limits to avoid deterministic effects and a single career limit (of 4 Sv) based on a doubling of the cancer risk in men aged 35 to 55. Later, when risk estimates for cancer had increased and were recognized to be age and sex dependent, the NCRP, in Report No. 98 in 1989, recommended a range of career limits based on age and sex from 1 to 4 Sv. NCRP is again in the process of revising recommendations for astronaut exposure, partly because risk estimates have increased further and partly to recognize trends in limiting radiation exposure occupationally on the ground. The result of these considerations is likely to be similar short-term limits for deterministic effects but modified career limits.
One major obstacle to human space exploration is the possible limitations imposed by the adverse effects of long-term exposure to the space environment. Even before human space flight began, the potentially brief exposure of astronauts to the very intense random solar energetic particle events was of great concern. A new challenge appears in deep-space exploration from exposure to the low-intensity heavy-ion flux of the galactic cosmic rays since the missions are of long duration, and accumulated exposures can be high. Because cancer induction rates increase behind low to moderate thicknesses of aluminum shielding, according to available biological data on mammalian exposures to galactic cosmic ray-like ions, aluminum shield requirements for a Mars mission may be prohibitively expensive in terms of mission launch costs. Alternative materials for vehicle construction are under investigation to provide lightweight habitat structures with enhanced shielding properties. In the present paper, updated estimates for astronaut exposures on a Mars mission are presented and shielding properties of alternative materials are compared with aluminum.
The radiation environment in low-Earth orbital flights is complex. It is strongly influenced by altitude, orbital inclination, time within a given solar cycle, flight duration, and shielding configuration. At any specified shielded location, both primary and secondary particles generated by nuclear interactions of primary particles with spacecraft structure are present. In addition, there are atmospheric secondary albedo protons and neutrons. No single detector can adequately measure this complex radiation field, and measurements of very high linear energy transfer target fragmentation products are particularly difficult. Crew radiation exposure have exclusively been measured using passive thermoluminescent detectors (TLDs). The cosmonaut exposures on the Mir station, uncorrected for the TLD inefficiency and neutron contribution, have varied from a low of 2.43 cGy to a high of 8.70 cGy. These correspond to dose rates of 144 microGy d(-1) to 468 microGy d(-1). These are consistent with rates observed by the D2 ion-chamber. Using the rates measured by the D1 chamber, dose rates under 4 cm of water vary from about 60 microGy d-1 to about 350 microGy d(-1). There is variation of about a factor of two between the dose rates at various locations in the same module. There is also a variation of dose rates of about a factor two between various modules. The highest astronaut dose for a Shuttle flight (STS-82) was 3.205 cGy with a dose rate of 3,221 microGy d(-1). Neutron contribution could be 36 +/- 15% of the astronaut charged particle dose equivalent. East-West asymmetry of dose rate is significant for spacecrafts that fly in an fixed altitude, such as the International Space Station.
Our basic understanding of the biological responses to cosmic radiations comes in large part from an international series of ground-based laboratory studies, where accelerators have provided the source of representative charged particle radiations. Most of the experimental studies have been performed using acute exposures to a single radiation type at relatively high doses and dose rates. However, most exposures in flight occur from low doses of mixed radiation fields at low-dose rates. This paper provides a brief overview of existing pertinent clinical and biological radiation data and the limitations associated with data available from specific components of the radiation fields in airflight and space travel.
Human exposures to ionizing radiation have been vastly altered by developing technology in the last century. This has been most obvious in the development of radiation generating devices and the utilization of nuclear energy. But even air travel has had its impact on human exposure. Human exposure increases with advancing aircraft technology as a result of the higher operating altitudes reducing the protective cover provided by Earth's atmosphere from extraterrestrial radiations. This increase in operating altitudes is taken to a limit by human operations in space. Less obvious is the changing character of the radiations at higher altitudes. The associated health risks are less understood with increasing altitude due to the increasing complexity and new field components found in high-altitude and space operations.
It is well established that labyrinth and neck afferent information contributes to the regulation of somatomotor function during movement and changes in posture. There is also convincing evidence that the vestibular system participates in the modulation of sympathetic outflow and cardiovascular function during changes in posture, presumably to prevent orthostatic hypotension. However, the labyrinth organs do not provide any signals concerning body movements with respect to the head. In contrast, the neck receptors, particularly muscle spindles, are well located and suited to provide information about changes in body position with respect to the head and vestibular signals. Studies in the cat suggest that neck afferent information may modulate the vestibulosympathetic reflex responses to head-neck movements. There is some evidence in the cat to suggest involvement of low threshold mechanoreceptors. However, human studies do not indicate that low threshold mechanoreceptors in the neck modulate cardiovascular responses. The human studies are consistent with the studies in the cat in that they demonstrate the importance of otolith activation in mediating cardiovascular and sympathetic responses to changes in posture. This paper briefly reviews the current experimental evidence concerning the involvement of neck afferent information in the modulation of cardiovascular control during movement and changes in posture.
Growing evidence suggests a role for the vestibular system in regulation of autonomic outflow during postural adjustments. In the present paper we review evidence for the patterning of sympathetic nerve activity elicited by vestibular stimulation. In response to electrical activation of vestibular afferents, firing of sympathetic nerves located throughout the body is altered. However, activity of the renal nerve is most sensitive to vestibular inputs. In contrast, high-intensity simultaneous activation of cutaneous and muscle inputs elicits equivalent changes in firing of the renal, superior mesenteric and lumbar colonic nerves. Responses of muscle vasoconstrictor (MVC) efferents to vestibular stimulation are either inhibitory (Type I) or are comprised of a combination of excitation and inhibition (Type II). Interestingly, single MVC units located in the hindlimb exhibited predominantly Type I responses while those located in the forelimb and face exhibited Type II responses. Furthermore, brachial and femoral arterial blood flows were dissociated in response to vestibular stimulation, such that brachial vascular resistance increased while femoral resistance decreased. These studies demonstrate that vestibulosympathetic reflexes are patterned according to both the anatomical location and innervation target of a particular sympathetic nerve, and can lead to distinct changes in local blood flow.
Data collected in both human subjects and animal models indicate that the vestibular system influences the control of blood pressure. In animals, peripheral vestibular lesions diminish the capacity to rapidly and accurately make cardiovascular adjustments to changes in posture. Thus, one role of vestibulo-cardiovascular influences is to elicit changes in blood distribution in the body so that stable blood pressure is maintained during movement. However, deficits in correcting blood pressure following vestibular lesions diminish over time, and are less severe when non-labyrinthine sensory cues regarding body position in space are provided. These observations show that pathways that mediate vestibulo-sympathetic reflexes can be subject to plastic changes. This review considers the adaptive plasticity in cardiovascular responses elicited by the central vestibular system. Recent data indicate that the posterior cerebellar vermis may play an important role in adaptation of these responses, such that ablation of the posterior vermis impairs recovery of orthostatic tolerance following subsequent vestibular lesions. Furthermore, recent experiments suggest that non-labyrinthine inputs to the central vestibular system may be important in controlling blood pressure during movement, particularly following vestibular dysfunction. A number of sensory inputs appear to be integrated to produce cardiovascular adjustments during changes in posture. Although loss of any one of these inputs does not induce lability in blood pressure, it is likely that maximal blood pressure stability is achieved by the integration of a variety of sensory cues signaling body position in space.
The human health community has been slow to adopt remote sensing technology for research, surveillance, or control activities. This chapter presents a brief history of the National Aeronautics and Space Administration's experiences in the use of remotely sensed data for health applications, and explores some of the obstacles, both real and perceived, that have slowed the transfer of this technology to the health community. These obstacles include the lack of awareness, which must be overcome through outreach and proper training in remote sensing, and inadequate spatial, spectral and temporal data resolutions, which are being addressed as new sensor systems are launched and currently overlooked (and underutilized) sensors are newly discovered by the health community. A basic training outline is presented, along with general considerations for selecting training candidates. The chapter concludes with a brief discussion of some current and future sensors that show promise for health applications.
The existence of obvious morphological and physiological changes in living systems exposed to altered gravity immediately informs us that prior changes have taken place in the chemistry of exposed cells, tissues and organs. These changes include transients that return more or less promptly to the norm when the system is restored to the terrestrial g-field. For example, altered serum hormone and electrolyte levels in man, which appear to reflect successful adaptation to the conditions of orbital weightlessness, disappear shortly after return to Earth. Other changes--in mineral and protein constituents of the skeletal system in man, and cell wall composition in plants--are more persistent or even permanent. Hypogravitational departures from the norm include not only "weightlessness" as achieved in orbit, but also experimental modes of compensation, on the clinostat or by flotation. These techniques are useful in the study of hypogravity but cannot replace fully the weightless environment. Plant ethylene and peroxidase both increase under orbital, clinostat and/or flotation conditions whereas 3-phosphoglyceraldehyde-dehydrogenase increases under orbital but not clinostat conditions; cytochrome reductase and malic dehydrogenase levels are affected by the clinostat, but not by actual weightless conditions. How do the altered organismal biochemistries induced by the centrifuge and the clinostat relate to one another? Does gravity operate on living systems as a continuous variable from 0 to superterrestrial values, or do deviations from g(earth) generate non-uniform, discontinuous stress responses, irrespective of sign? In plants, measurements of wall lignin content and peroxidase activity yield opposite answers. Given the limited data so far available we will consider the meaning of these contradictions.
This review describes the renal-endocrine mechanisms related to the early losses of fluid-electrolytes from the body during weightlessness as well as their contribution to longer term adaptation of fluid-electrolyte balance. The hypotheses presented were generated by a systematic analysis of body fluid and renal dynamics observed under conditions of actual and simulated spaceflight. These have increased our understanding of the effects of acute headward fluid shifts on renal excretion, the factors promoting excess sodium excretion and the regulation of extracellular fluid composition.
The impact of life on the atmosphere is examined through a discussion of the budgets of important atmospheric constituents and the processes that control their concentrations. Life profoundly influences oxygen and a number of minor atmospheric constituents, but many important gases, including those with the greatest effect on global climate, appear to be little altered by biological processes, at least in the steady state.
The endosymbiotic hypothesis for the origin of the eukaryotic cell has been applied to the origin of the mitochondria and chloroplasts. However as has been pointed out by Mereschowsky in 1905, it should also be applied to the nucleus as well. If the nucleus, mitochondria and chloroplasts are endosymbionts, then it is likely that the organism that did the engulfing was not a DNA-based organism. In fact, it is useful to postulate that this organism was a primitive RNA-based organism. This hypothesis would explain the preponderance of RNA viruses found in eukaryotic cells. The centriole and basal body do not have a double membrane or DNA. Like all MTOCs (microtubule organising centres), they have a structural or morphic RNA implicated in their formation. This would argue for their origin in the early RNA-based organism rather than in an endosymbiotic event involving bacteria. Finally, the eukaryotic cell uses RNA in ways quite unlike bacteria, thus pointing to a greater emphasis of RNA in both control and structure in the cell. The origin of the eukaryotic cell may tell us why it rather than its prokaryotic relative evolved into the metazoans who are reading this paper.
Prochloron is a genus of prokaryotic algae with photosynthetic pigments like those of chlorophytes. Prochlorophytes are almost invariably found associated as symbionts with marine protochordates (didemnid ascidians), and so far none has been successfully grown in sustained culture away from in host. Based on materials collected from nature, information of various sorts (biochemical, physiological, cytological and fine-structural) has been obtained, indicating many resemblances (and probably close phylogenetic affinities) between prochlorophytes and cyanophytes. Nevertheless they are distinguished by certain unique combinations of characters. Some of the data support the symbiogenesis theory for the origin of green-plant chloroplasts. Other possibilities are briefly discussed.
Substance P inhibits renin secretion. This polypeptide is a transmitter in primary afferent neurons and is released from the peripheral as well as the central portions of these neurons. It is present in afferent nerves from the kidneys. Neuropeptide Y, which is a cotransmitter with norepinephrine and epinephrine, is found in sympathetic neurons that are closely associated with and presumably innervate the juxtagolmerular cells. Its effect on renin secretion is unknown, but it produces renal vasoconstriction and natriuresis. Vasoactive intestinal polypeptide (VIP) is a cotransmitter with acetylocholine in cholinergic neurons, and this polypeptide stimulates renin secretion. We cannot find any evidence for its occurence in neurons in the kidneys, but various stimuli increase plasma VIP to levels comparable to those produced by doses of exogenous VIP which stimulated renin secretion. Neostigmine increases plasma VIP and plasma renin activity, and the VIP appears to be responsible for the increase in renin secretion, since the increase is not blocked by renal denervation or propranolol. Stimulation of various areas in the brain produces sympathetically mediated increases in plasma renin activity associated with increases in blood pressure. However, there is pharmacological evidence that the renin response can be separated from the blood pressure response. In anaesthetized dogs, drugs that increase central serotonergic discharge increase renin secretion without increasing blood pressure. In rats, activation of sertonergic neurons in the dorsal raphe nucleus increases renin secretion by a pathway that projects from this nucleus to the ventral hypothalamus, and from there to the kidneys via the sympathetic nervous system. The serotonin releasing drug parachloramphetamine also increases plasma VIP, but VIP does not appear to be the primary mediator of the renin response. There is preliminary evidence that the serotonergic neurons in the dorsal raphe nucleus are part of the pathway by which psychosocial stimuli increase renin secretion.
Developments during the first ten minutes of geotropic stimulation in plant seedling shoots are reviewed. Topics include induction and curvature; early processes; the relationship between auxin, electric field, calcium, and differential growth; gravity reception leading to Went-Cholodny transport; and comparison of root and shoot. Early processes reviewed are sedimentation of amyloplasts, release of ethylene, rise of electrical and auxin asymmetry, redistribution of calcium, asymmetric vascular transport, increase in tendency to deposit callose, and simulation of putative exocytotic voltage transients.
The hypothesis that extinction events have recurred periodically over the last quarter billion years is greatly strengthened by new data on the stratigraphic ranges of marine animal genera. In the interval from the Permian to Recent, these data encompass some 13,000 generic extinctions, providing a more sensitive indicator of species-level extinctions than previously used familial data. Extinction time series computed from the generic data display nine strong peaks that are nearly uniformly spaced at 26 Ma intervals over the last 270 Ma. Most of these peaks correspond to extinction events recognized in more detailed, if limited, biostratigraphic studies. These new data weaken or negate most arguments against periodicity, which have involved criticisms of the taxonomic data base, sampling intervals, chronometric time scales, and statistical methods used in previous analyses. The criticisms are reviewed in some detail and various new calculations and simulations, including one assessing the effects of paraphyletic taxa, are presented. Although the new data strengthen the case for periodicity, they offer little new insight into the deriving mechanism behind the pattern. However, they do suggest that many of the periodic events may not have been catastrophic, occurring instead over several stratigraphic stages or substages.