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i

1

(NASA-CF.- 1 )54 36) THE ALPHA-HELIX CONCEPT: N 84- 2 0610

1NNOVAIIVE JEILI..AilUN OF !Hz SPACE STATIGN

VL(UG (AM. A aEPUHT iU TdE bATICNhi.

AERONAUTICAL AND SPACE ADMINISTRATION Unclas

hL,UESTING ES'TAdLISHMENT CE A (Michigal, Gs/ 15 00o46

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k ^ ,'^ 'ter

CMG t`\^ c^^>^ ,-'a

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THE ALPHA-HELIX CONCEPT

Innovative Utilization of the

Space Station Program

NASA Grant •-).: NASW 3748, ORD 33060

3/24/83 to 10/17/83, $ 18,294

Robert S. Bandurski
- THE ALPHA-1

Innovative Utilization o:

A report to the

and Space AI

Requ

Establishment of

Laboratory on

NASA Grant No.:

' S

i ..
ORIGINAL PAM IS

OF POOR QUALITY

Principal Investigator:

Robert S. Bandurski, Professor

Department of Botany and Plant Pathology

Michigan State University

East Lansing, MI 48824-1312

517/355-4685

Co-Investigator:

ec lays.,

Dr. Neelam Singh

Department of Botany and Plant Pathology

Michigan State University

East Lansing, MI 48824-1312

517/355-6589

Department Chairperson:

Edward J. Klos, qairman

Department of any and Plant Pathology

Michigan State University

East Lansing, MI 48824-1312

517/355-:680

Contract and Grants:

Mr. E. B. Stauffer

Contracts and Grants

or

H. G. Grider, Director

Contract and Grants Administration
2

THE ALPHA-HELIX CONCEPT

ABSTRACT

It is proposed that a major laboratory - dedicated to biological-medical

research - be established on the Space Platform. The laboratory would focus on

Sensory Physiology and Biochemistry since Sensory Physiology represents the

first impact of the new space environment on living organisms. The objective

would be to use "microgravity" and the "high radiation" environment of space to

help solve the problems of prolonged sojourns in space but, more importantly, to

help solve terrestrial problems of human health and agricultural productivity.

The emphasis would be on experimental use of microorganisms and small plants and

small animals to minimize the space and time required to use the Space Platform

for maximum human betterment.

In the following document we introduce the "Alpha Helix Concept", that is,

the use of the Space Platform to bring experimental bio-medicine to a new and

extreme frontier - so as to better understand our worldly environment. We propose staffing and instrumenting the Space Platform Bio-Medical laboratory in a

manner patterned after successful terrestrial sensory physiology laboratories.

PART I: INTRODUCTION

How does life survive at the extreme environments of our world? That,

question represents the Alpha-Helix Concept. How does life cope and persist at

the temperature extremes of the world where enzymatic rates drop precipitously

or at the other extreme, where enzymes are literally cooked. How does life

persist where salt water meets fresh and survive an osmotic challenge or where

aerobic and hyperbaric conditions exist and life survives by bringing into play

new metabolic pathways. This use of extremes,. to better understand our mesic
conditions, is the Alpha-Helix Concept and was the life dedication of the late

Professor Per M. Scholander of the Scripps Oceanographic Laboratory.

Scolander's dedication resulted in the Alpha-Helix, a floating, earth

bound, biochemistry laboratory. This ship brought new meaning to many

bio-medical areas. Now we can look again at the Alpha-Helix concept - not with

the limitations of our worldly environment in mind - but with the expanded and

new dimensions of space. All life evolved in a one

advent of the Space Platform, we can, for the first

gamut of life processes in a "micro g" environment.

organelle has "weight". How has nature used weight

of life? We wish to answer that question.

What of radiation? All life has evolved under

moisture and carbon dioxide. But what are the effe,

"q" environment. With the

time, examine the entire

Every cell and cell

to accomplish the objectives

a protective screen of

:ts of solar radiation? Has

it acted as a positive evolutionary force or placed us in danger of extinction?

We wish to answer that question.

These are the dimensions presented by this new Alpha-Helix - a Space

Platform that will, for the first time, enable man to study and understand the

impact of gravity and solar radiation on life processes. No one can say where

this study will lead. But there is one certainty: just as temperature, and

salinity, just as elements such as oxygen, nitrogen, phosphorus, sulfur, etc.,

shaped our existence so will we find that gravity and radiation have shaped our

existence. Gra-.ity and radiation may prove to be environmental hazards - or

they may prove to be life givi p q forces - and now these questions can be

answered.

t
4

The Plan:

But, how to capitalize on this new opportunity? It is so new that no plan

is certain. But there is an old-fashioned homology that can serve as a guide -

"if it works - don't fix it", and so this report suggests again using the plan

which served so well on earth. To wit - build a space platform - Alpha Helixl

Just as the Alpha Helix carried the best instrumentation of its time to the

frontiers of the world - the new Space Platform should carry a very sophisticated physiological-tiochemistry laboratory to this new frontier. We can carry

what is now known of the instrumentation of sensory physiology and biochemistry

into space. It is towards this end that the body of this r-aport deals with

examples of sensory physiology and biochemistry. Sensory physiology is stressed

because it is the first impact in the long cascade of responses of an organism

to a stimulus. How do organisms sense light and gravity? How do they transduce

this perception into a life-affecting response? What is the molecular basis of

environmental perception? Wh , t machinery do we require to answer these

questions?

The Alpha-Helix Laboratory:

To exactly plan this laboratory will require the combined efforts of the

NASA laboratories; the Life Science-Space Biology Group; other physico-chemical

scientists and other workers in sensory physiology. Certain needs can be

outlined as follows:

Instrumentation:

First, it is fortunate that Fourier transform - Infra Red Spectrometry;

High Field Nuclear Magnetic Resonance Spectrometry; Tandem Mass Spectrometry;

Electron Diffraction Microscopy; capillary gas chromatography and high

performance liquid chromatography and computer control of all of the above have

developed - just in time. Now, for the first time, physical instrumentation is

adequate for the micromolar region of the biological world. No one individual

^
5

could specify the instrumentation required, but an example of how this could be

done is furnished by Dr. Robert Rabson, of the Department of Energy, who has

recently convened a group under the leadership of Dr. Elijah Romanoff of the

National Science Foundation to look at the interface between bio-medicine and

physico-chemical instrumentation. An extension of this approach might serve as

a model.

This report suggests convening our best physico-chemical instrumentalists

together with our best bio-medical sensory experimentalists to exactly specify

the sensitivities and capabilities of the instrumentation which musL, be carried

on the space platform to determine how living organisms sense gravity and

radiation and transduce the perception of these stimuli.

Physical Dimensions of the Alpha Helix Laboratory:

` Three things are required: First, space for the necessary instrumentation,

secondly, a micro "g" environment immediately adjacent to a variable, zero to

one "g" centrifuge, and thirdly, a radiation shielded environment, immediately

adjacent to an unshielded environment. Space should be allocated to small

plants and animals - and probably - for these first adventures we must emphasize

creatures which are very small and rapidly complete their life cycles - such as

- fungi, nematodes, bacteria, mice and small plants.

Earth based experimentation:

No amount of space studies have relevancy unless they are based on the data

provided by terrestrial studies such as provided by the Life Sciences Division

of the Space Biology Program. The question must always be - What are we sensing

in comparison to earth-based environments? That is the take home question.

The scope of this report:

The bulk of this report deals with the minutae of our knowledge of the

physiology and biochemistry of sensory physiology. The detailed conclusions are
not vital to the purposes of this report, - what is important is that we can use

earth-based studies, the Alpha-Helix Concept, to show what physico-chemical

tnstrumentation has helped solve terrestrial problems and how such instrumentation will solve new problems in space biology. In short, we propose using

"earth bound" sensory physiology and biochemistry as a guide to the new

Alpha-Helix laboratories.

PART II: MOLECULAR COMPONENTS OF THE ENVIRONMENT SENSING SYSTEM

The Bacterial Chemoreceptor System

Introduction

Most of our knowledge of this system has arisen from studies of two enteric

bacteria, Escherichia coli and Salmonella typhimurium (1,2). Studies of these

gram-negative bacteria have demonstrated such similar systems that the data are

. regarded as interchangeable. Further, studies on a gram-positive bacterium,

such as Bacillus subtilis, and even on photosynthetic bacteria confirm that

similar systems are operative even in these distantly related species. Also, as

will be discussed below similarities between bacterial chemotaxis and higher

plant and animal behavior are so striking as to indicate that we are dealing

with life processes that are common to all living things. Thus, there is a

clear advantage to using the reductionist approach to study the simplest things

first.

Now to briefly describe the system. The bacterium has slender thread-like

appendages scattered over the cell surface. When these cilia- appendages -

beat in an uncoordinated manner the bacterium tumbles in a random, Brownianmotion-type manner. The bacterium is just as likely to tumble away from a food

source as towards it. Now comes the magic! If the bacterium encounters a

dissolved nutrient then the cilia fuse into an oar-like flagellum and the

bacterium swims in a straight line. If the nutrient concentration increases -

:"

_
indicating the bacterium is swimming towards the nutrient source - the bacterium

continues to swim in that straight-line direction. If, however, the nutrient 

concentration decreases then the flagella dissociate into dispersed cilia and

th3 bacterium resumes random tumbling. Thus, moving away from the nutrient

source causes random tumbling, whereas, if by chance the bacterium starts on the

right course - it continues on that course.

The analogy to things even so complex as human behavior is obvious. We

tumble until we reach the right course and then that behavior is reinforced and

we move in the correct direction. Even memory is involved since the bacterium

must "remember" whether the nutrient concentration is increasing or decreasing.

.'ie following description of microbial chemozaxis is given in the hope that

a) it will guide us in laboratory design, and b) guide us in studies of more

complex systems.

The Sensory Receptor

First, the organism must perceive the environment. The best understood

environment receptors are the chemoreceptors of the enteric bacteria - that is

the receptors which recognize a favorable chemical and - ultimately - enable

the bacterium to swim towards that nutrient chemical and/or away from toxic

substances. Thus, most of this section will be devoted to the kinds of

experimentation and equipment which led to our present knowledge of bacterial

chemoreceptors.

Many other kinds of receptors have, however, been studied. A few examplep

are receptors for: light, sound, heat, temperature, touch, moisture,

osmolarity, and gravity. Unfortunately gravity perception is among the input

perceptions that is least well understood and as to radiation, other than a

narrow band in the visual region, our knowledge is very limited. 
6

Nonetheless, as will be seen from these few examples there is a certain

pattern to the experimentation and one can try to seek out experimental patterns

and something of the equipment and facilities needed.

The bacterial chemoreceptor:

A bacterium must swim towards a favorable nutrient and away from a toxic

chemical. To do so requires, of course, the motility apparatus, but also a

means of receiving the chemical signal - and, most importantly - a signal memory

device. The bacterium must measure the nutrient concentration at point "A" and

"B" and remember whether A is larger or smaller than B. This is the only way a

bacterium can swim towards a nutrient and away from a toxicant.

The bacterial chemoreceptor

To date, all chemoreceptors have been found to be proteins. They are

highly specific, usually recognizing only 1 or 2 compounds, and they have a high

affinity for that particular compound with a dissociation constant (K p ) of the

order of 10- 7 M. For example the galactose-glucose receptor of S. typhimurium

has a Kp for galactose of 2x10 -7 M, for glucose, 10 -7 M, for arabinose, 10-5

M, for lactose, 10 -4 M, for fucose, 10- 3 M and no binding for rioose. The

specificity and affinity of a chemoreceptor protein is thus comparable to that

of an enzyme and, indeed, the similarity to an enzyme is great except that

enzymes catalyze chemical changes in their substrate whereas chemoreceptors,

usually, release their substrate unchanged.

The specificity of chemoreceptors can be studied very easily in vivo simply

by placing a capillary pipette filled with the attractant into a beaker

containing the bacteria with or without the competitive attractant. If there

is no competition, the bacteria concentrate in the capillary. If there is

competition, the bacteria remain distributed in the beaker.

For detailed studies, however, the chemoreceptor must be isolated, purified

to homogeneity - if possible - and then Kp's be determined using equilibrium

dialysis or related techniques.
9

Once the purified chemoreceptor is avvilable it also becomes possible to

determine the amino acid composition and hopefully ultimately to even determine

tertiary structure of the folded protein. Thus, with effort, we can "see" the

pocket in the protein into which the chemical fits.

This then must become a major objective - to identify the environmental

acceptors and to see if there is a commonality of amino acid composition in the

composition of these receptors.

Analogies to other systems:

The analogies to the chemotactic system of bacteria are the photo and

gravity-responsive systems of higher plants and animals. For example, if in

plants and animals the stimulus of light oz gravity caused the release of a

chemical messenger, and this chemical messenger then came to rest on a

chemoreceptor - much as in the bacterial chemoreceptor system - then there would

emerge an exact analogy between bacterial chemotropism and higher organism behavior and a unity of ho:: organisms respond to their environment would emerge.

The important point is that a study of environment-receptor proteins must

be planned for on the space platform.

Photoreceptors for fungi and higher plants (4,5):

The best studied photoreceptor is that of the fungus Phycomryces blaksleeanus

and even those studies are based primarily on action spectra - that is - the

response of the organism to light is measured as a function o` wavelength.

Obviously action spectra can be misleading owing to distorting factors such as a

possible dependency of quantum yield on wavelength and, most importantly, the

presence of screening pigments. Nonetheless, for both Phycomyces and higher

plants there is reasonable certainty that the absorbing pigment is either a

flavin or carotenoid.

^

-^
10

Despite the uncertainties concerning the photoreceptor pigment, it is

possible, as pointed out above and by Delbruck, that we deal with a special case

of chemoreception - "the chemical in question being a photochemical product"

(4). Delbruck believed that the amount of photoproduct would be proportional to

the number of incident quanta and their cross-sectional capture area. Cross

sectional capture area should be proportional to the molecular extinction

coefficient of the light capturing pigment. Thus, the flux of quanta at each

wavelength to give a standard effect should provide an absorption spectrum of

the photoreceptive pigment. Similar reasoning has been applied to many photo

effects in higher plants.

Here again we introduce a positive note. Possibly the photo- and geo-act

releases a product which binds to a receptor - and thus chemo, photo- and

geo-perception are exactly analogous. The point is that once the receptor and

its effector are united - they can depolarize a membrane and initiate all the

processes of growth.

It is obvious that there is an analogy between terrestrial chemoreception

of bacteri-, and photoreception of plants and fungi and the problem of

georeception by higher organisms, plants and microorganisms.

The action spectra (efficacy as function of wavelength) was measured by

Delbruck and Shropshire (5) by interposing on a Phycomyces sporangiophore - an

upright, hair-like strand of the fungus - a standard light of broad wavelen gth a

test light of variable and narrow wavelength. In this manner they then could

measure the bending of the sporangiophore and they found an action spectrum with

clear maximum at 485, 455, 385 and 280 nm. This action spectrum fits the

absorption spectrum of a flavin - although, unfortunately, it could also be a

carotenoid. Thus we must leave this terrestrial problem with the ambiguous
I 

conclusion that for fungi, as well as higher plants, there is uncertainty as to

whether the photoreceptor for blue 'light is a Flavin or carotenoid.

The georeceptor:

The problem of the georeceptor introduces even more uncertainties. The

prevalent notion is that some heavy cytoplasmic body falls to the base of the

cell and there - in an unspecified manner - initiates appropriate growth

responses such that a vertical orientation is again attained (6,7).

The problem is that thorn are no real controls. That a cellular microbody

denser than the surrounding cytoplasm falls to the bottom of the cell is no

surprise. But is this any more a message of gravity than Newton's falling

apple? Indeed, Newton could have concluded that the apple tree sensed the

direction of gravity by the direction of the fall of the apple.

• Clearly, this is a problem for the fipace platform laboratory. We have

definite and refined knowledge of the chemoreceptors of bac*eLia. These are

characterized and sequenced proteins with a high affinity for the appropriate

chemotactic chemical.

We have no comparable knowledge for the photo and gravity reception

systems. Does the light or gravi , .y release a chemical that then binds to a

"chemoreceptor" protein - or do light and gravity circumvent this initial step

of chemoreceptor complex and go immediately to a transduction signal for the

appropriate biological response?

This is a major problem of terrestrial - as well as space biology. How do

organisms transdtice an environmental signal into an appropriate biological

response?
12 

Phycomyces sensor mutants that are deficient in both photo and geo responses

(3).

Many mutants of Phycomyces with abnormal phototropism have been prepared

and studied. Of the greatest interest is the fact that these mutants are very

often also deficient in their response to gravity as well as response to other

tropic stimuli.

Briefly, the system is as follows: Phycomyces has a giant spore-bearing

aerial hyphae called a sporangiophore which will readily attain a height of 10

or more centimeters. The sporangiosphore is really one giant multi-nucleated

cell with one central vacuole. It shows the following tropic responses: First

it is phototropically sensitive and will resoond to a stimulus of about 500

quanta (Shropshire) of 440 nm light; second, it responds to gravity and will

grow again into an upright position if placed horizontally; thirdly, it can be

photoinduced by blue light to synthesize beta-carotene; and lastly, and most

strikingly, it evidences a phenomenon called "autochemotropism". Autochemotropism means that the sporangiophore will grow away from an object placed in

its growth path, for example, a glass rod, without even touching the obstructing

object. How this avoidance phenomenon is accomplished is completely veiled in

my stery. It has been postulated that the sporangiophore releases a gas -

identity unknown - and reflectance of the gas causes the avoidance response.

Nonetheless there are then at least the 4 types of tropic responses and these

can be studied in mutant strains.

Mutagenesis was accomplished with nitrosoguanidine and then a most

ingenious method was used to select for the mutants. The mutated spores were

grown in closed containers illuminated with a horizontal beam of light. Normal,

non-mutant sporangiophores do not grow horizontally towards the light because of

the gravity response ar: iastead grow at an angle of 30 0 . This 30 0 angle is
j

OF POOR QUALITY

attained over a factor of 10 9 variation in the intensity of light. Thss the

r '

i sporangiophore is attracted to the light but cannot grow directly towards the

I

i light owing to the corrective gravity response which would cause the sporangiophore to grow vertically upright. Thus the sporangiophores could be scored for

i their phototropic response, their geotropic response; also their beta - carotene

content and their autochemotropic response. An example of the autochemotropic

response of the wild type ( NRRL1555) and two mutant strains is shown in the 

figure below ( taken from Bergman et al., p. 10).

I

n 0.1 1 10

700

m 100

so

r

dark -s 0 •=

1092 1

I 

By this means it was possible to arrange the mutants into 3 phenoL^ypic

classes as shown in the figure below (8).

georeceDlor 

memoreteplor

IOSSII IQiS1 CIOSS2

C21 C33 C6

C47 C39 C63

photo- U09 C,4 UPIroDUms 

rttepror rill C69 0107 

0112 C94 Cu0

0114 CO 0149

^nduc l ion of

0-corolene

synth! S's

inihofron of

sporo,q,oDhores
ORIGINAL PAGE M 14

^)F POOR QUALITY.

The remarkable thing is that the mutants do not fall into the full 16

classes expected if the 4 criteria were independent. Instead they fall into 3

general classes.

These remarkable experiments again indicate that somehow environmental

receptors are plugged into only a few transduction systems which then give the

appropriate growth response. This is a strong argument towards using a "micro

g" and high radiation environtnent to better learn how small organisms perceive

and then transduce an environmental stimulus. Knowledge gained from these

experimentally advantageous organisms will certainly have general applicability

to mammalian physiology.

T!-.e tropic systems of plants

Plants respond both to light and to gravity. Further, as to light, they

measure the length of the light (and, or dark) photoperiod and the wavelength of

the light. These light responses have been thoroughly reviewed elsewhere (9).

Concerning the gravity sensing component of plants - there is much less

certainty (7,8). As shown below, the plant can begin to respond to gravity

within minutes - probably instantaneously (10).

Q09 TRANSDUCER ARM i

Y

COLEOPTILE 

_ QO COLEOPTILE I MESOCOTYL 

it

NODE

a QO

v

• 003-

Q01 -"MESOCOTYL

0 2 a a 8 10

TIME (minutes)
15

The time period for the response seems so short as to preclude the settling of a

heavy organelle to the bottom of the plant cell. Further, in the case of young

plant shoots, the shoot responds uniformly - to the gravitational stimulus along

its entire length - governed only by its intrinsic growth rate.

Thus, for gravity, we do not know what the gravity sensing system is.

There is much literation concerning "statoliths" and, indeed, there are even

studies of mutants and of starved plants indicating that "starch-free" plants no

longer respond to gravity. Thus a case can be made indicating that the plant

senses gravity by means of a heavy, starch-like organelle. Still, the time

course of the response - and the similarity to other tropic responses seems to

argue against this simple mechanical explanation.

Conclusions concerning receptors:

Any laboratory dealing with receptors for gravity and for radiation sensing

elements must be equipped as would a modern protein chemistry laboratory on

earth. There is no way one can "freeze" a protein in its intricate tertitary

configuration, and bring it back to earth for detailed study. The "Alpha Helix"

Space Platform laboratory must contain the accoutrements of an excellent protein

structure laboratory - as defined by current practitioners of the art.

The equipment required would include that necessary for sequencing both

proteins an.; nucleic acids together with the ancilliary equipment required for

preparing the pure proteins.

PART III: TRANSDUCTION OF THE RECEPTOR RESPONSE (2,11,12)

Once the stimulus is perceived - whether it be light falling on a

photoreceptor, gravity perceived by some unknown apparatus, or a chemical

attractant combining with an appropriate receptor site - there arises the

problem of the transduction of this (receptor-stimulus) complex to some

meaningful physiological response.

s. -
16

Again, it is the bacterial chemoreceptor system that is best understood.

Mutations in three genes, abolish chemotaxis (8). The bacterium is still 

actively motile but it doesn't swim towards an attractant or away from a toxic

substance. It can still bind the attractant molecule. Thus, it is believed 

that thesc transducer proteins are transmembrane proteins that relay information

to the flagella concerning the degree of occupancy of the chemoreceptors (Boyd &

Simon) .

Somehow the receptor protein reacts with the attractant - with a certain

rate constant and as a function of the concentration of the attractant. There

is also a rate constant for the rate of dissociation of receptor and attractant.

It is this rate of formation and rate of breakdown of [receptor-attractant] that 

governs the degree of receptor occupancy. Thus the transducer protein must

"tell" the flagellar apparatus what the degree of occupancy of the receptor is.

This then represents the simplest form of memory - the transducer protein

measures at two times, t 1 and t2 , how many receptors are occupied. One

cannot help but wonder whether human sort term memory - in one form or another

- also measures "occupancy" of a neurotransmitter site.

The nature of the signal from transducer protein to the flagellar motor

apparatus -_prolonged effects:

The transducer then has 3 domains, an external one which measures receptor

occupancy, an internal domain where chemical changes in the transducer must

occur and lastly, a domain where the transducer causes disperse cillia to form

into a swimming-effective, flagella.

What changes occur in the transducer? There appears to be an immediate

effect followed by a more gradual prolonged effect (2). The prolonged effect

involves transducer methylation and sensory adaptation is an overall measure of

the activities of a methyltransferase and a methyl esterase. A certain level of
17

stimulus can be correlated with a certain steady-state level of methylation of

the transducer (2). Thus, when the organism is moving toward the attractant,

there is a block to methyl esterase activity, thus causing a transient rise in

the level of methylated transducers.

Of the very rapid short range effects we know much less. In higher plants

(13), in the case of mammalian neurotransmitters (14), in the case of ciliated

protozoa (11) as well as in the enteric bacteria one sees rapid changes in

membrane potential.. Possibly the occupied receptor protein opens a gated pore

which changes the cellular Ca++ , Na + and K+ concentrations.

How the stimualted receptor protein transfers its knowledge to a transducer

and then how that transducer affects a motor apparatus represent weak points

in our knowledge. These points, however, are absolutely central to any

understanding of behavioral psychology and of how the environment "instructs"

both plants and animals to behave. The over-all importance of this knowledge to

our terrestrial well-being cannot be overemphasized.

PART IV: Summary and specific recommendations for a sensory physiology

laboratory on the space platform:

There is attached to this a small list of references which concern specific

statements in this report. In addition, there is attached a list of references

thot will serve as an introduction to these aspects of sensory physiology.

References to mammals, in general, and human physiology, in particular are left

to other reports in this program. Here we have focused only upon a few of the

best studied microbial, protozoan and seedling plant systems.

We regret these omissions and restrictions. The field of sensory

physiology is so hugh as to require a multi-disciplinary approach by many

workers. With that caveat in mind, we make the following recommendations:

^
1) It is apparent that the medical problems of space physiology are closely

related to the problems of sensory physiology in lower plants, animals and

bacteria.

2) It is clear that the response to an environmental signal, terrestrial or

"micro-g", is mediated by: a) the signal; b) a receptor protein; c) a

transducer and; d) a response apparatus.

3) There is great similarity in the manner in which sensory physiology research

is conducted, including, a) use of mutant organisms; b) the techniques of

protein and nucleic acid isolation and characterization; c) the methods of

small molecule chemical characterization; d) electrophysiological techniques

and, e) microscopy.

4) As stated in the Introduction, there is need for side by side comparisons of

a "micro-g" environmental with the environment of a 0 to 1 g centrifuge.

5) There is need for a heavily radiation shielded environment - preferably a

"layered" environment adjacent to a region with minimal shielding.

6) To the best of our current knowledge - based on terrestrial studies of

sensory physiology - it would be best to equip the laboratory exactly as one 

would equip a sophisticated chemistry-biochemistry laboratory. The equipment

would include: a 0°C working area - that is a small "walk-in" cold room; the

centrifuges and chromatography and electrophoresis apparatus required for

protein and nucleic acid research; a sterile hood and chemical fume hood; a

wet lab area; and the standard IR, NMR, and mass spectral equipment.

7) Owing to the complexities of the above described studies, it would be best to

use teams of perhaps 3 scientists with orientations in biology, genetics and

chemistry-biochemistry.
19

PART V. A Short List of Selected References

1) Koshland, D. E., Bacterial chemotaxis as a model behavioral system, Society

of General Physiologists, Volume, Raven Press, New York ( 1980).

2) Boyd, A., and Simon, M., Bacterial Chemotaxis, Ann. Rev. Physiol., 44,

501-17 (1982).

3) Glazer, A. N., Comparative biochemistry of photosynthetic light-'harvestinq

systems, ann. Rev. Biochem. 52, 125 -57 (1983).

4) Bergman, K., Burke, K. V., Cerdo -Olmedo, E., David, C. N., Delbruck, M.,

Foster, K. W., Goodell, E. W., Heisenberg, M., Mussuer, G., Zalokav, M.,

Dennison, D. S., and Shropshire, W., Bacteriol. Rev. 33, 99-157 ( 1969).

5) Shropshire, W., Stimulus Perception, in Encyclop. of Plant Physiol., Ed. by

W. Haupt and M. E. Feincleib, 7, 10-41 ( 1979), Springer-Verlag, Berlin.

6) Sievers, A., and Hensel, W., Gravitropism, In: Plant Growth Substances,

1982, P. F. Wareing, Ed., Academic Press, London ( 1982).

7) Bandurski, R. S., and Schulze, A., Gravitational effects on plant growth

hormone concentration, Adv. Space Res. 3, 229-235 (1983).

8) Bergman, K., Eslava, A. P., and Cerda-Olmedo, E., Mol. Gen. Genetics 123,

1-16 (1973).

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21

PART VI: CHEMOTAXIS

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PART VII: PHYCOMYCES

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of Phycomyces. II. Distribution of stretch and twist along the growing

zone, and the distribution of their responses to a periodic illumination

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toxic responses to the stimulation of narrow test areas. J. Gen. Physiol.

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Proc. Natl. Acad. Sci. U.S.A. 73:1969-1973.
52

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53

Jayaram, M., D. Presti and M. Delbruck. 1979. Light induced carotene synthei is

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57

PART VIII: GRAVITY AND GEOTROPISM (PLANT)

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PART IX: PHOTOTAXIS AND PHOTOTROPISM

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