Engineering topics
Publications and source records attributed to Brown, A. H..
Research in plant physiology consists mostly of studies on plant growth because almost everything a plant does is done by growing. Most aspects of plant growth are strongly influenced by the earth's gravity vector. Research on those phenomena address scientific questions specifically about how plants use gravity to guide their growth processes.
Experiments were undertaken to determine if the reciprocity rule is valid for gravitropic responses of oat coleoptiles in the acceleration region below 1 g. The rule predicts that the gravitropic response should be proportional to the product of the applied acceleration and the stimulation time. Seedlings were cultivated on 1 g centrifuges and transferred to test centrifuges to apply a transverse g-stimulation. Since responses occurred in microgravity, the uncertainties about the validity of clinostat simulation of weightlessness was avoided. Plants at two stages of coleoptile development were tested. Plant responses were obtained using time-lapse video recordings that were analyzed after the flight. Stimulus intensities and durations were varied and ranged from 0.1 to 1.0 g and from 2 to 130 min, respectively. For threshold g-doses the reciprocity rule was obeyed. The threshold dose was of the order of 55 g s and 120 g s, respectively, for two groups of plants investigated. Reciprocity was studied also at bending responses which are from just above the detectable level to about 10 degrees. The validity of the rule could not be confirmed for higher g-doses, chiefly because the data were more variable. It was investigated whether the uniformity of the overall response data increased when the gravitropic dose was defined as (gm x t) with m-values different from unity. This was not the case and the reciprocity concept is, therefore, valid also in the hypogravity region. The concept of gravitropic dose, the product of the transverse acceleration and the stimulation time, is also well-defined in the acceleration region studied. With the same hardware, tests were done on earth where responses occurred on clinostats. The results did not contradict the reciprocity rule but scatter in the data was large.
We conducted a series of gravitropic experiments on Avena coleoptiles in the weightlessness environment of Spacelab. The purpose was to test the threshold stimulus, reciprocity rule and autotropic reactions to a range of g-force stimulations of different intensities and durations The tests avoided the potentially complicating effects of earth's gravity and the interference from clinostat ambiguities. Using slow-speed centrifuges, coleoptiles received transversal accelerations in the hypogravity range between 0.l and 1.0 g over periods that ranged from 2 to 130 min. All responses that occurred in weightlessness were compared to clinostat experiments on earth using the same apparatus. Characteristic gravitropistic response patterns of Atuena were not substantially different from those observed in ground-based experiments. Gravitropic presentation times were extrapolated. The threshold at 1.0 g was less than 1 min (shortest stimulation time 2 min), in agreement with values obtained on the ground. The least stimulus tested, 0.1 g for 130 min, produced a significant response. Therefore the absolute threshold for a gravitropic response is less than 0.1 g.
Dark-grown wheat coleoptiles developed strong curvatures within 5 h of being transferred in orbit from a 1 g centrifuge to microgravity during an experiment flown on the IML-1 shuttle mission. The curving tendency was strongest in seedlings that were immature, with coleoptiles shorter than 10 mm at the time of transfer. The curvature direction was non-random, and directed away from the caryopsis (the coleptile face adjacent to the caryopsis becoming convex). The curvatures were most marked in the basal third of the coleoptiles, contrasting with phototropic responses, which occur in the apical third. We interpret these curvatures as being nastic, and related to the curvatures commonly reported to occur during clinostat rotation treatments.
In January 1992, the NASA Suttle mission STS 42 carried a facility designed to perform experiments on plant gravi- and photo-tropic responses. This equipment, the Gravitational Plant Physiology Facility (GPPF) was made up of a number of interconnected units mounted within a Spacelab double rack. The details of these units and the plant growth containers designed for use in GPPF are described. The equipment functioned well during the mission and returned a substantial body of time-lapse video data on plant responses to tropistic stimuli under conditions of orbital microgravity. GPPF is maintained by NASA Ames Research Center, and is flight qualifiable for future spacelab missions.
The principal objective of the research reported here was to determine whether a plant's periodic growth oscillations, called circumnutations, would persist in the absence of a significant gravitational or inertial force. The definitive experiment was made possible by access to the condition of protracted near weightlessness in an earth satellite. The experiment, performed during the first flight of Spacelab on the National Aeronautics and Space Administration shuttle, Columbia, in November and December, 1983, tested a biophysical model, proposed in 1967, that might account for circumnutation as a gravity-dependent growth response. However, circumnutations were observed in microgravity. They continued for many hours without stimulation by a significant g-force. Therefore, neither a gravitational nor an inertial g-force was an absolute requirement for initiation [correction of initation] or continuation of circumnutation. On average, circumnutation was significantly more vigorous in satellite orbit than on earth-based clinostats. Therefore, at least for sunflower (Helianthus annuus L.) circumnutation, clinostatting is not the functional equivalent of weightlessness.
This time course and location of gravitropically induced curvatures in stems of goosegrass (Gallium aparine L.), a member of the Rubiaceae, have been investigated. In the early stages of the response (0-5 h), curvature develops throughout the growing region, and is followed by an autotropic straightening which affects the internodes only, leading to the production of essentially straight internodes some 15 h after the onset of gravistimulation. Curvatures developing in the nodal regions, however, continue to increase over this period, and are not subject to reversal by autotropism. The nodal curvatures are not entirely dependent on the presence of any other part of the plant, since marked curvatures can be induced in isolated nodal segments. This pattern of response leads ultimately to correction of the growth direction of the plant by means of curvature responses confined exclusively to the nodes, despite the initial participation of both nodes and internodes in the gravitropic reaction.
The phototropic dose-response relationship has been determined for Triticum aestivum cv. Broom coleoptiles growing on a purpose-built clinostat apparatus providing gravity compensation by rotation about a horizontal axis at 2 rev min-1. These data are compared with data sets obtained with the clinostat axis vertical and stationary, as a 1 g control, and rotating vertically to examine clinostat effects other than gravity compensation. Triticum at 1 g follows the well-established pattern of other cereal coleoptiles with a first positive curvature at low doses, followed by an indifferent response region, and a second positive response at progressively increasing doses. However, these response regions lie at higher dose levels than reported for Avena. There is no significant difference between the responses observed with the clinostat axis vertical in the rotating and stationary modes, but gravity compensation by horizontal rotation increases the magnitude of first and second positive curvatures some threefold at 100 min after stimulation. The indifferent response is replaced by a significant curvature towards the light source, but remains apparent as a reduced curvature response at these dose levels.
Circumnutations of hypocotyls of sunflower (Helianthus annuus L. cv. Californicus) were studied under 1 g and 3 g conditions. Root mean square values of the hypocotyl deviation from the plumbline and period of the movements were determined from calculations of the autocorrelation functions of the movements. The amplitude and the period of the circumnutations increased under 3 g as compared to 1 g. A transition from 3 to 1 g or vice versa also caused changes in period and amplitude of the movements. The results are interpreted as a support for the idea that gravity influences the circumnutation parameters in this sunflower variety. A comparison is made with published results on the dwarf sunflower cv. Teddy Bear where the force influence is very small or negligible. Simulations of a model for circumnutations show movements which are in qualitative agreement with the experimental results, provided adaptation to g-levels is included in the model. Finally, the results are discussed with the recent Spacelab-experiment (SL1) as a background.
Flight experiments on the importance of gravity to plant growth and behavior are reported. The following studies were undertaken: (1) hyperastic responses to incremental changes of an axially imposed centripetal force; (2) Spacelab-1 experiments, methods for preparing soil in flight hardware containers were impound, to ensure desired moisture content and minimal contamination probability; (3) mesocotyl growth patterns were established by Avena lore exposure to red light during early seedling outogency; (4) the development of flight hardware; (5) choice of member of seedlings in each cube; (6) data processing and reduction; (7) clinostat validation; circummutation in space was more vigorous than on Earth based clinostat.
We report a pioneering attempt to use the NASA Shuttle Orbiter Middeck locker facility to acquire data on plant growth in near weightlessness. The information was needed to confirm the suitability of a plant culture system to be used in an experiment scheduled for the first Spacelab mission. The test was designed to measure germination and early seedling growth in a series of soil mixtures covering a range of water contents. Empirical determination of growth dependence on moisture content was required because both in theory and from Soviet flight experience it seemed possible that the dependence function in near weightlessness could be critically different from what we had measured on Earth. Such a difference could invalidate the future test in Spacelab 1 of gravity dependence of the differential growth process, circumnutation. After two failed attempts sufficient measurements were obtained from the third Shuttle Orbiter flight test to confirm the biocompatibility of the plant culture system--viz. soil moisture content variations had the same effect in near weightlessness as at 1 g. A number of supplemental observations about middeck locker conditions in Shuttle flight are presented. These may prove helpful to would-be experimenters who will plan to take advantage of future Shuttle flight opportunities for biological research.
For over half a century and especially since the 1960's a number of plant physiologists, seeking to explain the impressively ubiquitous mechanism that drives and regulates circumnutation in all growing plant organs, have been unable to agree on whether the differential growth process that leads to circumnutational oscillations is gravity dependent. There has been fairly general agreement that the question might be answered, if test plants could be deprived of all significant gravitational stimuli as would be possible in the near weightlessness or free fall environment of satellite orbit. Such an experiment was carried out during the Spacelab 1 mission. Circumnutational oscillations were observed which demonstrated that a protracted input of gravitational information from the environment was not required for initiation or maintenance of circumnutation in sunflower hypocotyls.
The requirements to and capabilities of a Space Station biological facility centrifuge are discussed on the basis of an assessment of the objectives and subjects of future microgravity biological experiments. It is argued that the facility should be capable of both acute and extended chronic exposure of test subjects and biological materials to altered-g loading. In addition, the experimental approaches and equipment for microgravity studies on a Space Station are outlined. Finally, the engineering requirements of such a centrifuge are examined, with consideration of radial gravity gradients, size, and physical access to animals.
Accomplishments during the past fiscal year consisted of (1) completion of research on a study of the kinetics of damping out of circummutation when the axially directed g-force was abruptly eliminated (Research Task NULYRL). (2) Further experiments were accomplished on a comprehensive study, underway for several years, to validate (or invalidate) the use of clinostat rotation as a hypogravity simulation device. (3) Some of our earlier (unpublished) observations on plant seedlings resistance to g-loading were evaluated, interpreted, and a paper was submitted for publication (Research Task HYGEFF). (4) In what has been called ""Shuttle middeck locker ecology" an attempt to acquire reliable empirical information on the thermal profile experienced by test packages housed in middeck lockers (MDDL) during shuttle flights at long last seems to be yielding some results.
Arabidopsis plants were grown in centrifuge tubes under well standardized culture conditions. Each plant was subjected to centrifugation (roots out) for 10 min at one of a series of centripetal forces between 7 and 390g. No deformation was observed in plants centrifuged at less than 35g. An 'average' degree of deformation was attained at about 60g. All plants exposed to more than 95g were maximally deformed but none was broken nor otherwise damaged irreversibly even at 390g. In every case new shoot growth continued normally after the centrifugation. A plant population grown on horizontal clinostats (0.5 rpm) under culture conditions exactly the same as for the upright plants responded to centrifugation stress in a way that was not substantially different from the response pattern of the plants cultured upright at 1g.
Ground based and spaceborne (Biosatellite) research into the effects of hypogravity on biological systems is summarized. A concept of sensitivity to gravity is defined: sensitivity = dR/dQ x 1/standard deviation of R, where R is the measured biological response to a quantitative stimulus, Q. Detection and transformation of gravity information by the organism are discussed. The role of an uncommon gravity force environment as stimulus for physiologic adaptations, rather than stress, is introduced.
The methods and results are presented concerning experiments using the first plants to make a journey for scientific purposes in earth orbit on board Space Shuttles 2 and 3. These experiments were conducted in order to validate the culture system planned for use in a Spacelab-1 experiment involving the growth and kinetics of sunflower seedlings. The experimental flight package is described and the preparation steps taken prior to the flights are discussed. The conditions to which the plants were exposed during the flights are examined in detail, focusing on the temperature profile and the gravitational forces during reentry to landing.