d,_.

o, SP ,_-14 _,

ON TI IE HABITABILITY
C,_SE F ILEOF MARS
approaCc_ _anetary ecosynthesis tP o

ONAL AERONAUTICS AND SPACEADMINISTRATION

_.
US_

..,_..,

NASA

SP-414

ON TI IE HABITABILITY OF MARS
unapproach to planetaryecosynthesis
Edited by M. M. Averner R. D. Mac EIroy

Prepared by Ames Research Center

Scientific and Technical Information NATIONAL AERONAUTICS

Office AND

1976 SPACE ADMINISTRATION Washington, D. C

CONTRIBUTORS R. D. MacElroy Program Ames Research Coordinator Moffett Field, CA Center,

M. M. Averner Dept. of Biology, Southern Oregon College, Ashland, OR

S. Berman Earth Sciences Dept., State University W. R. Kuhn Dept. of Atmos. and Oceanic Sci., Univ. of Michigan, Ann Arbor, MI College, Oneonta, NY

P. W. Langhoff Dept. of Chemistry, Indiana Univ., Bloomington, IN

S. R. Rogers Dept. of Biological Science, Dartmouth College, Hanover, NH

J. W. Thomas Biological Sciences Dept., Cal. Poly. State Univ., San Luis Obispo, CA

For sale by the National Technical Information Service Springfield, Virginia. 22161 Price - $5.25

PREFACE This study of the Faculty authored Planetary Fellowship contributions was conducted Biology Program which and at NASA-Ames with the NASA. Research Center, under the auspices

Division

financial

support

of the Stanford-Ames of individually by the editors.

The report

is a compilation and integrated

have been assembled

iii

ACKNOWLEDGMENTS The Lederberg proddings development and by Hans Ichtiaque of this study Mark. Later, Rasool was stimulated extensive crystallized the by conversations with James concept. Harold with Joshua Danielli, Klein, and Donald

discussions

DeVincenzi and Richard Young were Before and during the study, discussions and Richard participation, sciences. Edward Joel ularly Lederberg,

responsible for making with Martin Alexander,

the study possible. Edward Leadbetter,

Vondrak focussed certain problems. Joel Levine's enthusiasm, willing and criticism carried us over many hurdles encountered in the physical people Richard Von to have contributed Bruce Orgel, Leslie their ideas and Anbar, work has criticism Whitten, Robert Billingham. to the James Rein, study: Pollack, Joshua Vondrak, White, Robert

Many Griffith, John

Levine, indebted

Eshelman, Carl

Michael

Oro, J. R. Garrels, Sagan,

H. Lettau,

and John

We are particexamined to all these

whose

published

previously

several of the ideas incorporated in this study. We are very grateful people for helping us into the beginnings of Planetary Engineering.

R. D. MacElroy

TABLE

OF

CONTENTS Page

CONTRIBUTORS PREFACE I. SUMMARY 2. REPORT PRESENT Limits Survival ACKNOWLEDGMENTS DIGEST

....................... ..................... ..................... CONDITIONS ............. .............. CONSIDERATIONS ..................... ..................... Organisms on Mars: ENVIRONMENT ..... ................. MARTIAN

ii iii v 1 3 3 4 4 5 6 7 7 ......... 8 9 10 11 OF MARS INVENTORY CLIMATOLOGY .............. ................ Solar Constant ....... ................ ................ Surface ..... ORGANISMS .......... .......... ......... 13 13 17 19 26 27 29 29 33 35 ................. 35 BIOTA ......... ...... ...... 38 43 45

.......................... ........................ MARTIAN to Life

BIOLOGICAL Photosynthesis

of Terrestrial

Computer Simulations MODIFICATION OF THE Greenhouse Effects

...................

Advective Heating .................... PROSPECTS FOR GENETIC ENGINEERING CONCLUSIONS ..................... 3. STUDY APPROACH .................... 4. PHYSICAL CHARACTERISTICS PHYSICAL AND CHEMICAL SURFACE SURFACE Spectral TEMPERATURE ENERGY Distribution RADIATION Considerations Transmission BUDGETS of the Martian

ULTRAVIOLET Geometrical Atmospheric

Ultraviolet Intensity Estimates for the Martian 5. THE TRANSPLANTATION OF TERRESTRIAL TO MARS A MARTIAN LICHENS ........................ ECOLOGY AS POSSIBLE MARTIAN

CYANOPHYTES AS POSSIBLE MARTIAN BIOTA BIOLOGICAL CYCLING OF ELEMENTS ON MARS

vii

6. SURVIVAL AND PHOTOSYNTHESIS F TERRESTRIAL O ORGANISMS ON MARS: COMPUTER MODELSFOR LICHENSAND CYANOPHYTES ............... GENERAL CONSIDERATIONS ...............
Organism Resistance Carbon LICHEN Temperature .................. Photosynthesis .....................

51
52 52 52 53 53 53 56

to Water Transpiration and Dioxide Diffusion ................ ................... ................ MODIFICATION ................. SURFACE TEMPERATURE ..... ................ OF THE

SCHEMATIC

CYANOPHYTE SCHEMATIC RESULTS ........................ 7. PLANETARY MARTIAN INCREASING ADVECTIVE ENGINEERING: ENVIRONMENT MARTIAN TRANSPORT

63 64 69 75 75 ......... OF .......... ......... ........ GROWTH 85 85 87 90 90 92 93 93 ...... 93 95 95 96 99 101 78 81 ........

8. GENETIC ENGINEERING .................. STRATEGY ....................... PROSPECTS 9. CONCLUSIONS 10. APPENDIX A: OF LICHENS GENERAL GENERAL Resistances Cyanophyte Plasmalemma Cytoplasmic FOR GENETIC ENGINEERING ...................... BIOLOGICAL CHARACTERISTICS AND CYANOPHYTES OF MODEL ON MARS ............. MODEL MATHEMATICAL ASPECTS APPROACH

PARAMETERS

OF CYANOPHYTE

to Water Transpiration ............. Mat - Resistances to CO2 Diffusion Resistance Resistance to CO2 Diffusion to CO2 Diffusion .......... ...........

Estimation of Substrate Constants for Photosynthesis PARAMETERS OF LICHEN MODEL ............ Resistance to Water Vapor Diffusion ............. ............ ............ Parameters for Lichen Photosynthesis 11. APPENDIX B: GREENHOUSE EFFECT 12. REFERENCES .......................

viii

TABLES Number 1 COMPARISON OF LIMITS TO LIFE WITH SELECTED MARTIAN ENVIRONMENTAL PARAMETERS ....... BIOLOGICAL SOME ORGANISM PHYSICAL (a) (b) (c) (d) (e) Planetary AND GROWTH CHARACTERISTICS AND AN OF IDEAL 6 OF ............... [% by volume] Incident ......... EARTH ............ AND MARS 13 13 14 14 14 at the Surface 15 Incident ............... K ............ ............. ON FOR MARS AND 18 ON 21 ON 22 ................. (g/cm 2) at the 15 15 16 16 Page 4

TERRESTRIAL

ORGANISMS

...................... CHARACTERISTICS and Orbital Parameters Parameters Composition ................... ................... Radiation Temperature,

Atmospheric Atmospheric Surface Average Winds

Solar Radiation

(cal/cm 2/day) (f) Average Ultraviolet Surface (g) Average (i) Surface

(cal/cm 2/day)

(h) Polar Cap Parameters A Chemical Inventory

DIURNAL HOURS LATITUDES ESTIMATION OBSERVED ESTIMATION

TEMPERATURE PER DAY AND ABOVE SEASONS

EXTREMES FREEZING ..............

SELECTED

OF K FOR EARTH, RADIATION FLUXES OF K FOR MARS,

BASED ............ BASED

THEORETICAL ANNUAL EARTH ANNUAL FOR

RADIATION ENERGY

MODEL

........... FOR

SURFACE AND MARS

BUDGETS

.................. TEMPERATURES CLIMATONOMY SURFACE ................. AND ........... RANGES

23

SURFACE FROM DIURNAL FROM ATMOSPHERE

MARS

25

MARTIAN VARIATIONS A DRY

TEMPERATURE ASSUMING 26

CLIMATONOMY,

ix

10

SOLAR

VACUUM

ULTRAVIOLET .................... SPECTRAL

SPECTRAL 28 DISTRIBUTION .... 28

DISTRIBUTION 11 12 SOLAR

ULTRAVIOLET OZONE

MOLECULAR CROSS

PHOTOABSORPTION ................... MOLECULAR ............... OZONE TRANSMISSION 31 PHOTOABSORPTION 32 AT THE MARTIAN 33 TO LIFE WITH SELECTED ....... FOR 44 VARIOUS ........ FOR ICE 66 CORRESPONDING 69 PRESSURE, FLUX OF AN SOME IDEAL 77 POLAR ........ 72 35 OZONE 31 30

SECTION

13

VACUUM

ULTRAVIOLET FUNCTION

TRANSMISSION 14

ULTRAVIOLET MOLECULAR FUNCTION ...................... MOLECULAR CROSS CARBON

15

DIOXIDE

SECTION

...................

16

ULTRAVIOLET INTENSITY SURFACE ...................... COMPARISON MARTIAN OF LIMITS

17

ENVIRONMENTAL ENVIRONMENTAL OF BLUE-GREEN

PARAMETERS EXTREMES ALGAE ...........

18

OBSERVED GROWTH

19

OXYGEN PRODUCTION BY LICHEN FOR LICHEN THICKNESS AND WIND SPEEDS TEMPERATURE, EQUILIBRIUM AND VAPOR PRESSURE, AND SLOPE

62

2O

BETWEEN CARBON ..................... PRESSURE, AND

DIOXIDE

21

TEMPERATURE, SLOPE OF SURFACE

ICE-WATER VAPOR ...................... BETWEEN AND AIR

SUBLIMATION

22

RELATIONSHIP TEMPERATURE,

ADVECTIVE

23

BIOLOGICAL

CHARACTERISTICS

TERRESTRIAL ORGANISMS AND MARTIAN ORGANISM .................

FIGURES Number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 Logic diagram Diurnal surface at equinox of study temperature ..................... ......... .................. for Mars; at the equator 17 20 46 46 and 49 54 ....... 55 57 58 ........... ........ ....... rate for lichen ............. 60 60 61 66 67 68 polar cap ..... 70 83 rate for lichen Page 12

Schematic energy budget for Martian surface The biological nitrogen cycle ............... The biological sulfur cycle ................ Biological recycling sulfur on Mars Lichen model Algae mat model Diurnal variation Diurnal algae Diurnal Diurnal Diurnal Carbon variation variation variation variation dioxide of carbon, nitrogen, .................... .................

schematic

schematic ................ of blue-green algae temperature of transpiration rate for blue-green

........................ of lichen temperature of transpiration of photosynthetic greenhouse effect

Pressure-temperature surface for carbon Water vapor greenhouse Average Proposed annual energy strategy

cross section of the thermodynamic dioxide and water ........... effect ............... for north of the Martian flux diagram

for the alteration .....................

atmosphere

xi

1. SUMMARY

The examined. state, and requirements several still

possibility Available chemical and

of utilizing data, inventory environmental the

Mars

as a habitat and are of

for terrestrial speculations and on life. water of the an man.

life, the

including climate, with accurate the

man, physical known data

is

assumptions, of Mars limits amount

reviewed terrestrial

compared While and

on are

points, no

particularly fundamental,

of Martian limitation lack of

nitrogen of Mars

reserves,

lacking,

insuperable The

ability

to support atmosphere

a terrestrial would surface oxygen of however, by (1)the by by upon strains climate rely prevent

ecology the

is identified. unaided is an habitation additional

oxygen-containing The The may present creation be feasible such might strong of

of Mars by major on time of years.

ultraviolet an adequate the use

irradiation and

barrier. Mars to period

ozone-containing organisms. be several of of of

atmosphere The millions novel, planetary the

through

photosynthetic might

needed This and caps

generate

an atmosphere, reduced Martian would and the

be drastically the present

synthesis techniques techniques the melting

Mars-adapted, engineering, engineering. polar the polar over Martian

oxygen Such and

producing climatic

photosynthetic

of genetic

(2) modifying

modification advective would period. require

concomitant however, long

greenhouse investment

heating of very

effects. large

Melting

caps,

amounts

of energy

a relatively

2. REPORT PRESENT

DIGEST MARTIAN CONDITIONS in the solar system variation of sunlight the Martian is about environment is closest to that

Of all the planets of Earth. The diurnal

the same on Mars as on Earth,

although the intensity of solar radiation at the surface of Mars is only about 60% of that at Earth's surface. The Martian day is 24.6 hr, and the year and seasons are twice Earth as long as those and is primarily provides of Earth. carbon Ozone The atmosphere dioxide. is present no shielding Oxygen is only comprises 1/200 as dense as that of only 0.1% or less of the and, other than at radiation of water which at the on or in the studies may be

mass of the atmosphere. the poles, equator averages Of critical planet. indicate regolith, The that other

in very small amounts to solar ultraviolet is the 0.1% amount

essentially

7 10 a ergs/cm 2/sec. importance to habitability contains suggest less than that only may be as much

atmosphere there estimates

water

vapor.

While recent this amount

as 1019 kg of water

ice in the polar caps and the water is available on Mars

1%, or less, of

present. The assumption is made in this study that enough to support the growth of terrestrial types of organisms. The average temperature of the planet is about However, in low- and mid-latitudes the temperature 7 hr a day in the summer. 300 K, at night winds generate unsheltered The Martian Martian on Mars, studies dust-carrying organism. surface ecology. surface temperature Temperature were calculated developed is, of course, ranges very important While the daytime their speed that could it falls to a low of 170 K. Although indicate storms could cause

60 K lower than that of Earth. rises above freezing for about may reach a maximum is known mechanical about stress 100 m/sec. of little surface to an of a

temperature approach severe

Such winds

to any discussion heating cycles

for annual at the

and diurnal University

for the in which

by using the method Lettau

of "climatonomy," of Wisconsin

a quantita-

tive approach

by H.H.

energy fluxes are expressed as Fourier time series. The mean annual temperature difference from summer to winter is calculated to be about 45 K in mid-and

polar-latitudes, 100 K at the when allows

and equator the

half

this

value and

at the

equator.

The

diurnal

range except

is slightly during This

over winter tech-

all year, range

is the same the

for mid-latitudes, the diurnal that terrestrial the

solstice nique 270

is 20 K. At of the limit number

poles

range

is zero.

estimation lower hr per

of hours of most

surface

temperature This

exceeds number from is the

K, a typical 2000

for growth year

organisms. planet

roughly equator

Martin

in a region

around

extending

to 45 S. CONSIDERATIONS

BIOLOGICAL

Limits

to Life Life can exist only and in certain the chemical must environments. constituents be obtained and Such of soil, factors water, as temperature, and air impose not light, limits

water on for life;

availability, a certain factor.

minimum The ability of the

a certain

maximum

exceeded by

each

of an environment environment 1. with

to support the known

life can be determined limits to life. A compari-

comparing son for Mars

parameters is provided 1.-

in table

TABLE

COMPARISON OF LIMITS TO LIFE WITH ENVIRONMENTAL PARAMETERS Range compatible Growth: 255 K-377 with life

SELECTED

MARTIAN

Parameter Temperature Ultraviolet irradiation

Mars 140 K-300 K

Survival: <79 K-377

3 l0 s ergs/cm 2 0 - 100% 10% - 100% (w/w) 0% - 35% NaCI 0-13 Abrasive particles _> cell size will cause mechanical damage

7 103 ergs/cm 2/sec 20 Present, amount

9

Oxygen (p02) Liquid H2 0 Salinity pH Mechanical abrasion

unknown

? Dust in storms _ 10-50/IM in size range of many microorganisms Not yet found Not yet found Not yet found CO, CO2, 5 mbar H20
r

Nitrogen Sulfur Phosphorus Carbon Oxygen Hydrogen

Absolute

requirement

H_O

When those allow growth

factors

whose hostile simple

values

can be measured not dependent Ultraviolet on the Martian range which

or estimated upon light, oxygen, particularly

are considered, oxygen would the anaerobes; in the range to their mean

Mars is an environment generally, single-celled

to biology. organisms.

The lack of atmospheric

only of those is lethal.

organisms Organisms

of 2000A-3000A,

surface would

would be exposed decrease

a mean flux of ultraviolet light in this survival times to a few minutes at most. The Martian surface temperature 7 hr a day at the equator and mid-latitudes.

range

would

allow seasonal

growth

for only

The low temperatures

and atmospheric

pressures would allow surface water of the planet, and the total amount chemical elements

to exist in a liquid state only in restricted areas of water can only be guessed. Other important and phosphorus, have yet to be identithe kinds of terrestrial are quite limited, and There to simulated

for life, such as nitrogen

fied on Mars. Even a most optimistic appraisal suggests that organisms able to survive in the present Martian environment the growth have been of even these many attempts forms would be quite restricted to determine the response

in vigor and extent.

of microorganisms

Martian environments. experimenters' choice

While conclusions have varied, in part a reflection of the of organisms and environmental conditions, investigators feel for growth of certain anaerobic, cold-adapted

that there is a definite possibility terrestrial bacteria on Mars. Photosynthesis For higher life forms,

an oxygen-containing

environment

is mandatory.

The

need for an oxygen atmosphere derives from two functions of oxygen: it is required environmentally for the formation of ozone which on Earth provides an ultravioletabsorbing organisms in principle, water eral consensus shield which under which life exists, dependent gaseous and biologically upon oxygen oxygen for metabolism for this purpose. in higher There are, from are absolutely oxygen-containing is that the present

two ways by which

can be produced: or biologically, atmospheric

chemically, oxygen

or other

compounds,

from water. The genwas biologiatmosphere was is expected to to make Mars and water of

mass of terrestrial

cally produced and the bulk of the oxygen in the ancient primitive produced by green plant or algal photosynthesis. Thus photosynthesis have the capability habitable. Photosynthesis into a carbohydrate of generating oxygen in the amounts fixing necessary

can be considered (starch)

as the

of carbon organisms

dioxide

by chlorophyll-containing

in the presence

visible light. As a by-product of photosynthesis, produced. In principle, all the necessary physical elements trial are available on Mars. The question organisms might be best then photosynthetic

oxygen, derived from water, is (light) and chemical (CO2, Hz O) is: which of the available terresto survive, grow and generate

fitted

oxygen Martian valleys those gests shown

on

Mars?

Although it is likely major ideal,

no that

Earth the

environment Antarctic groups. and dry

exactly valleys

resembles come their Martian are closest.

available In these with sugas

eco-niches, there required that none in table are four by are 2.

photosynthetic "ideal" lichens

Comparing algae

properties organism the "best"

a hypothetical although

oxygen-evoloving blue-green

All gen. times ever, tent All

groups groups

except except on

the the the

blue-green lichens are would

algae

have

an absolute to ultraviolet short. might of growth The be

requirement light, so that

for

oxy-

sensitive be and slow quite

survival howfull

unprotected can with occupy the goal

surface

blue-green shielded of lichens

algae, from is not the

substirface light. of rapid

eco-niches The very oxygen

thus rate

intensity

of ultraviolet

consis-

evolution. OF SOME ORGANISM

TABLE 2.- BIOLOGICAL AND GROWTH CHARACTERISTICS TERRESTRIAL ORGANISMS AND OF AN IDEAL MARTIAN Extreme resistance to ultraviolet radiation

Organism

Requires oxygen

Extreme resistance to drying

Growth rate

Growth habitat

Green algae

Yes

No

No

Fast (hr)

Soil (surface and subsurface), Snow (surface), water Surfaces (rock, tree)

Lichen Moss Blue-green algae Ideal Martian organism

Yes

Yes

Yes

Very slow (yr) Slow (wk) Fast (hr) Very fast (min)

Yes NO No

No No Yes

No Yes

Moist surfaces Soil (surface and subsurface), water Soil (surface and subsurface), water

Yes

Survival

of Terrestrial a biological with

Organisms standpoint

on Mars: several need organism

Computer questions vary during atmospheric

Simulations concerning For a diurnal the cycle, interaction how and would how How low of an the closely severe

From organism temperature would would and it

its environment

to be examined.

example,

of a hypothetical be coupled water to the

ground from

and the

temperatures? water,

be the strong

loss resulting characteristic

lack of atmospheric Can desiccation

pressures, limited by

winds

of Mars'?

be significantly

varying within likely

the

resistance limits? and

properties can estimates terrestrial genetic

of the these

structural various

components environmental production

of the be

organism affect Is it Martian

reasonable

How might

parameters made?

photosynthesis, conditions

of photosynthetic organisms engineering? could

that any present-day without extensive

survive the envisioned

Taking into consideration to water movement, ultraviolet formulated organism candidates models ature to yield estimates temperature. for growth

such parameters as photosynthetic rates, resistance sensitivity, and the like, computer models were of such factors as water flux, oxygen production, and two organisms which might be algae and a mat of lichens. The coupled with to the surface increasing wind temperspeed

These models incorporated on Mars: a mat of blue-green temperature would be closely

predict:

1. The organism's

rather than the atmospheric temperature. 2. The organism's temperature would of convective heat loss.

decrease

because

3. A layer of desiccated cells on the upper surface of the algal mat would control water loss. The lack of such a layer would permit large water losses from the lichen. The losses would, wind speed. 4. Temperature 5 hr/day. generate in the Allowing an amount Martian and in turn, be affected water loss would by such factors limit as lichen thickness to only of carbon yr. To and 3 to

photosynthesis of Mars, blue-green amount in 7000

a 25% coverage of oxygen

of the

surface

algae could dioxide an produce

equivalent (approximately

to the present 5 mbar)

atmosphere

amount equivalent to the minimum necessary for human breathing (approximately 100 mbar) would take 140,000 yr. The lichens would take approximately 10 times longer. MODIFICATION Greenhouse Many Effects of the environmental factors which are unfavorable for the establishment temperature. The OF THE MARTIAN ENVIRONMENT

of life on Mars would

be ameliorated

by modifying

the planet's

surface temperatures of Mars could be increased, and the diurnal temperature variations reduced, if a means could be found for increasing the atmospheric mass by vaporizing not known, the polar however, caps. The mass and composition the portion dioxide, the summertime of the sublimate cap remnant in the caps is grows may be carbon and although of the caps which seasonally

and recedes may be carbon dioxide, water, or both.

Increasing the amount of carbon dioxide or water vapor in the atmosphere will increase the surface temperature. This warming, known as "the greenhouse effect,"

occurs some the

because of the

certain

gases, radiation to space,

carbon (heat)

dioxide from

and

water

vapor

among

them, not and of north

absorb all of heats

thermal is lost

the surface down

of the from due to

planet

so that

radiation

but is re-emitted

the the

atmosphere injection remnant of carbon by

the surface. The amounts of carbon The cap might the were greenhouse entirely

increase dioxide effect carbon only pressure provides be added average which

in surface and water from dioxide, increase would

temperature was calculated. dioxide the mean some

various polar it

carbon then the increase larger

is small.

If the

maximum surface 100-fold.

amount temperature

dioxide

contain

would

7 K, although

atmospheric Water vapor could resulting water by vapor, The

a much to the be

greenhouse rise to effect would the the

effect; be

for

example, pressure If the

if enough by only caps and apprevapor

water 10%, contain carried ciable

vapor the

atmosphere released where greenhouse of sublimate heating than until the carbon for the

to increase

the total 10 K.

temperature could

remnant

ice then

atmosphere atmosphere be induced. has also the

as water could been

advection possibility

to lower a substantial of initial

latitudes greenhouse

support

could effect so

a "runaway" injection and

investigated: that This be howfor be

that

is,

could

an

increase

temperature occur? latter could

continued appears maintained ever, sufficient difficult. less

sublimation likely over time the for

greenhouse vapor pole of

would carbon

spontaneously dioxide. rose in the The several global

water

summer a new

temperature dioxide steady-state

degrees; atmosphere would

continued

maintenance to allow

temperature

to be established

Advective An atmospheric tudes tures climatic season. regime pressure pressure ways: outside 20%. from 0.25) such the by

Heating additional mass while factor increases, The mean equatorial which the global must amount temperatures given winter To times. in the the be considered is advective from change polar of 190 but there heating; tow are as the latistable winter stable air surface of from cap by reduced (albedo If

of heat

transported does not In brief,

to high two the

increases. increase regimes The exists at

temperature a carbon one attain This

polar

tempera-

decrease. dioxide stable

possible a polar

on Mars, surface 1 bar. ten bound

cap during The regime about K and

present

climate

represents

of the this

regimes.

second the

temperature high

at a surface

of approximately must be increased gases or 0.77 the by

temperature

might

be brought by importing solar polar admixture albedo by flux cap

in a number

volatilizing the planet, last present over solar

regolith, effective if the small cap's

an atmosphere over of sand the polar were or dust

increasing could be

This spread

possibility cap

realized the

albedo

to 0.73. would

A relatively lower were

the required

amount. years.

increased

absorbance

maintained

for approximately

a hundred

advective instability couldbe triggeredanda newhigh temperaturelimaticregime c established. Carbondioxide,althoughnot efficientin increasing globaltemperathe ture by a greenhouseffect may providepolar heatingby advection. olarheating e P couldcause injectionof increased mountsof watervaporinto the atmosphere, the a which would providefor a stronger reenhouseffect. Thus the combined g e mechanisms climate. of advective and greenhouse heating could be used to modify the Martian

PROSPECTS

FOR

GENETIC

ENG|NEERING

Modification heating would ing to would optimize speculate the

of increase

the

Martian under possibility

climate the area which of

by of

means the

of planet

advective available would species

and for occur. of

greenhouse growth and It is interest-

enormously

conditions the

such creating

growth novel

upon

photosynthetic Martian into environ"ideal"

organisms ment:

far better in effect, Such

adapted

to growth currently

in the

present "best utilizing

or modified fit"

transforming genetic

available

organisms

organisms. tion currently

engineering

is possible,

methods

of gene

manipula-

known

or under and apparatus

development. their viruses of cells. has yielded Genes for to the repair moving Several have species for the powerful the tools protein for the manipof cells. of of

Research ulation cells Thus and of the these

on bacteria genetic (to a great a certain

determine physical

enzymes repair

extent)

determine

characteristics of rapid gene lack

if a cell has radiation A cell Today them

characteristic, it is due such a rapid

example, presence genes of these been

capability of a certain would from donor techniques,

ultraviolet genes. genes. inserting

damage, lacks are recipient DNA

or of several this gene or and

which there into and

mechanism

techniques organisms.

organisms that

is, sexual of in

recombination blue-green characteristics several the different gene algae.

transformation, no ideal single an Martian

successfully algae well they

applied may might Indeed,

in species have be found

Thus defining species pool

while

of blue-green as gene available

all those

organism,

of algae of the Earth

all utilizable might be

donors. for the

in principle, of Continued synthesis genes of an

entire

construction

ideally advances genes then

adapted in our lead

oxygen-producing understanding to a potential into a recipient

photosynthetic of gene for "best structure novel fit"

Martian and of methods genes. These

organism. for the synthetic

may

creating

could

be integrated

organism.

CONCLUSIONS On the basis of currently 1. No fundamental, terrestrial life has been available information limitation identified. the study to the However, concludes important that: data are not

insuperable unequivocally

ability

of Mars to support

available, for example, the extent of water reserves and the composition of the polar caps. These data must be acquired before a more accurate assessment of the habitability of Mars can be made. tory. forms, carried injection be carried energy years. 4. Mechanisms could present terrestrial of genetic engineering currently available or under development be used to construct organisms. either the Martian environment or available photosynthetic organ5. Altering isms, or both, human habitat organisms far better adapted to grow on Mars than any 2. For human life to exist, the creation Such an atmosphere might be generated but the time required 3. Climate out of polar out equivalent modification cap sublimate by a combination may be in hundreds leading of advective of time of an oxygen atmosphere is mandaby present terrestrial photosynthetic of thousands in surface of years. temperature effects initiated would might be by the need to of

to an increase

and greenhouse at the total

into the atmosphere. of solar energy

This injection expenditure incident

for a long period

of amounts

to the total amount

on Mars over several

would significantly decrease the time required to create an acceptable on Mars. Indeed it may be mandatory to take these steps if Mars is to planet.

be made into a habitable

10

3. STUDY The decades

APPROACH technological advances which have been achieved planetary over the last three within there

leave little doubt

as to the feasibility

of future

exploration

our solar system.

While exploitation

of the planets

by man may seem remote,

is a distinct possibility that technological make the utilization of Martian resources

developments economically

or the needs of society may feasible or socially desirable.

To exploit Mars more efficiently it may be necessary to establish permanent human communities on the planet. The question thus arises as to whether Mars is a habitable planet or can exists be made into and Examples reserves, one. This has been have investigated reviewed would be: by our group. of Mars of Current whether being sufficient data, there inhabited water assumptions, by man. or nitrogen speculations insuperable of such an inability been to determine the absence

a fundamental,

limitation to increase

to the prospect intolerably

a limitation

low surface

temperatures, or the presence of toxic chemicals in the atmosphere or soil. Failure to identify such factors may only reflect the present lack of sufficient information. The continued acquisition necessary to clarify these habitability questions: 1. Can contemporary 2. If not, removed? terrestrial present what are of data on the physics and chemistry of Mars will be and other crucial aspects of habitability. To assess the participants posed and attempted to answer a series of Martian surface? constraints to the growth adapted be of

of Mars the study

terrestrial the climate

life grow on the present to growth and

constraints be modified

can these

3. Can the Martian orgahisms? 4. Can terrestrial or a modified

to be more conducive modified to be better

organisms Martian

be genetically

to the

environment?

These questions have been arranged in a flow chart (fig. 1). This particular logic is only one of many that might be considered and, often, a clear-cut yes or no decision is impossible at this time. The study group's answers to these questions are based upon speculation, mathematical models and data generated by extensive literature review, group discussion, and conversations with invited speakers (chs. 4 8). These answers have, in turn, led to a number 11 of conclusions (ch. 9).

PRESENT

WILL MARTIAN

NO PRESENT

WILL MARTIAN

NO BE

AN ORGANISM ENGINEERED IN PRESENT

N TO O

ENVIRONMENT

ENVIRONMENT

GROW

SOPPO.TE_MAN
yES r

SOPPORT
i YES

N_"%%'_&T
I YES

Wl HI LL T S WTH R GRO ESULT IN A PLANE] FIT FOR HUMAN

O NO

" GANISM

BE NO CAN BE TO MARS NO TO MAKE F T FOR MODIFIED HUMAN SUPPORT LIFE;'

WILL

MARS T HE A TO IF )_ NO

ENGINEERED GROW & PLANET

HAVE, IN F UTURE, POTENTIAL SUPPORT H MAN

HABITATION)

CAN AN HUMAN

YES

YES

YES

YES

i ,
I I ......... _ -I I

, ,_ , ISP_%_TST.ATEGY L_-SPECIFY 2 ECOLOGICAL CONSIOERATIONSL_ F 4 3 OTHER TIME I

Figure

1.

Logic diagram of study.

12

4.

PHYSICAL

CHARACTERISTICS

OF

MARS

PHYSICAL

AND

CHEMICAL

INVENTORY

To conditions major

assess on

the the

potential planet

of Mars is needed. and

for supporting Table surface 3 provides parameters

life,

an accurate

idea

of present of the The

an up-to-date for both both published Earth

summary and

planetary,

atmospheric,

Mars.

data have been extracted from a variety of sources, The references are indicated beneath each table. Table The led lack 3 reveals that Mars surface is a small, features. cold, These dry

and

unpublished.

planet

with and a rough,

a thin

atmosphere. effects has surface,

of an appreciable of harsh

atmosphere

with

its shielding include

moderating cratered

to a number

strong winds organizing flux at the ground, and

occasionally large diurnal

into global-scale duststorms, high ultraviolet temperature excursions of the order of 100 K

TABLE

3.-

PHYSICAL

CHARACTERISTICS

OF EARTH

AND MARS

(a) Planetary and Orbital Parameter Mass, g Mean density, g/cm 3 Mean radius, km Surface gravity, cm/sec 2 Length of day, Earth-days Length of year, Earth-days Obliquity, deg Orbital eccentricity Mean distance from Sun, km Solar constant, cal/cm 2/min Planetary albedo Effective temperature, K Source of data: Goody

Parameters Earth Mars

6.43X 1026

5.98X 1027 5.52 6371 981 1 365 23.5 0.017 150X 106 2.00 0.30-0.35 253

3.94 3394 373 1.026 687 23.9 0.093 228X 106 0.866 0.15 0.25 216

and Walker (1972). 13

TABLE 3.-

CONTINUED Parameters Earth 5.3X 102 t 1000 1.2X 10 -a 8.4 9.8 2 10 80 Mars 2.41019 5 1:2 10 -s 10.6 4.5 0.1 a 30? 150

(b) Atmospheric Parameter Atmospheric mass, g Surface air pressure, mbar Surface air density, g/cm a Scale height, km Adiabatic lapse rate, K/kin Average optical thickness Tropopause height, km Turbopause height, km Sources of data:

Goody and Walker (1972);

Noll and McElroy (1974).

aMuch higher in duststorms.

(c) Atmospheric Gas

N2

Composition Earth 78 20 1a 0.93 0.03 lO-S < 10-s with latitude (1971);

[% by volume] Mars
9

02 H20 Ar CO2 CO 03 aDisplays large variations

0.1 <0.1 a 1-25 99 98-74 0.1 < 6X 10-s a

and season. Noll and McElroy (1974);

Sources of data: Barth (1974);

McComlac

Levine (1975).

(d) Surface Winds Typical wind speeds 2 cm above the surface,mps 2 cm above the surface,raps Earth 0.5 5 Mars 5 50

Sources of data: Barth (1974); Noll and McElroy (1974); B. White (Univ. of Calif., Davis) (personal communication).

14

TABLE 3.(e) Average Solar Radiation Latitude, deg 90 45 0 45 90 alncludes N N S S attenuation

CONTINUED Incident at the Surface (cal/cm 2/day) Mars Northern Summer 320 315 250 100 0 turbidity Hemisphere Winter 0 100 365 450 450 and cloudiness.

Earth a Northern Summer 327? 520 380 99 0 Hemisphere Winter 0 112 419 513 341?

due to atmospheric

Sources of data: Unpublished notes of H. H. Lettau (Univ. of Wisconsin) (Earth values); Levine, Kraemer, and Kuhn (1974) (Martian values).

(f) Average Ultraviolet

Radiation

Incident

at the Surface (cal/cm2/day)

UV Band 2000-3000 A

Earth _ 0

Mars 10.6 a

aThis figure is equal to 6 103 erg/cm 2/sec. Source of data: Nawrocki and Papa (1963). Temperature, K Mars Northern Summer 185 220 200 162 145 Hemisphere Winter 145 175 240 265 200

(g) Average Surface Latitude, deg 90 N 45 N 0 45S 90 S

Eartha Northern Summer 279 289 297 279 226 Hemisphere Winter 235 261 297 287 263

Source of data: Crutcher (1969)(Earth values); Woiceshyn (1974); Conrath et al. (1973); Kliore et al. (1973); Hanel et al. (1972); (Mars values).

15

TABLE 3.-

CONCLUDED

(h) Polar Cap Parameters Earth Parameter NPC (Greenland) Latitudinal extent from pole: 1. Northern Hemisphere winter, 2. Southern Hemisphere winter, SPC (Antarctica) ] 60-90 85 -90 6.7 0.19 1? H2 O, CO2 ? 88-90 60-90 NPC Mars SPC

deg deg

...

o..

Percent of globe covered by cap: 1. Northern Hemisphere winter, % 2. Southern Hemisphere summer, % Average cap thickness, Composition: Sources of data: km:

2.7 1.7 1.5 H20

0.9 2.2 2.5 H20 (Mars).

0.03 6.7 0.017 H2 O, CO2 ?

Bates (1964)

(Earth);

Woicesnyn

(1974)

(i) A Chemical Species CO2 atmosphere: crust: H2 0 atmosphere: crust: 02 atmosphere: crust: N2 atmosphere: crust: Os atmosphere:

Inventory Earth 0.3 l0 s 2 106 200

10 7

(g/cm 2) Mars 15 10-10 a ? 0.01 1 103 .9 0.01

9

780 102?
10 -a

0.5 ? 1-4 .9 2X
10 -7

Sources of data: Barth (1974); Noll and McElroy (1974); Cannon (1974); Sagan (1971); Ingersoll (19741).

Fanale and

16

(fig. 2). The Martianpolar capsareintriguingsurfacefeatures. heyarebelieved bemostlywater T to

ice with limates winter. remnants speculated which tial this close needs greenhouse possibility, scrutiny Although water mates assumed have this huge been water vapor a thin carbon dioxide ice layer which subduring frozen been freeze substanof during summer The actual amount is unknown. to be only an The and redeposits of water in the polar caps in have deep

I 300

I _ /

GROUND

a: 200

atmosphere

to be thawed and advective polar regions

to initiate effects. have

Because come

100 0

I 2

I 4

I 6

I 8

I 10

I 12

I 14

I 16 hr

I 18

I 20

I 22

24

the

under

LOCAL

TIME,

in recent it exists is

years. generally Martian vary Levine the planet's Fanale's water agreed widely. (1975) that Based suggests The suggestion little estion Figure 2.- Diurnal surface temperature for Mars; at the equator at equinox. that as much 1975) at high Physical as 3.51023 happened there that g may to all may accounts to the altoon the be

in the

atmosphere,

of subsurface outgassing released remains

ratios, from lenses unanswered.

interior.

question (Fanale,

of what

underground 10-30% chemically no more reveals than

of ice or hard-frozen estimate water, percent. in current the fate of and The of water polar unsettled living

permafrost outgassed. cap remnants nature a gap

latitudes adsorption

for only surface, gether Mars

of Levine's bound a few gap

probably question

contain of water

of the that organisms

a serious project

knowledge,

is especially implanted

significant on

when trying to Martian surface.

terrestrial

SURFACE

TEMPERATURE

CLIMATOLOGY

Surface failure tively known narrow

temperature range, from

is one on

of

the

crucial 273

variables K to 373

in determining temperatures K, are detrimental

success

or

of terrestrial terrestrial It is important

organisms organisms. to note

Mars.

Environmental

outside

a relato most

approximately

that

surface

temperatures microorganisms,

are

of more role

concern in making with,

than

air Mars thus of the

temperatures, habitable controlled not the only thin

since is discussed by, varies the with

oxygen-generating more latitude There fully and in later

whose

sections, but has

will large

be in contact The surface variations diurnal

and

environmental

parameters season is also

at the surface. temperature

temperature because at

atmosphere. interface. insight from who to the figure developed

a significant

discontinuity

ground-air An be derived (1968),

magnitude 2 which a

of this discontinuity is constructed model 17 from of the the

and

the

diurnal of Gierasch lower

variations

can

data

and Goody atmosphere

theoretical

Martian

consistent ith the groundbased w observations f Martiansurfacetemperaturesy o b SintonandStrong(1960) andGifford (1956).Note that the air temperature variation is about80 K, the maximum temperature beingreached several hoursafterlocal noon,andthe minimumin the earlymorning.The surfacetemperature variationis evenlarger,beingsome 140 K. The largesttemperature discontinuityoccursat aboutlocal noon,the surface temperature eingsome70 K higherthanthe adjacent b air temperature. he sizeof the discontinuitydepends T directlyon the amountof solarradiationreaching surface. hus,at latitude 45 N, duringthe winter,the the T differencebetween andsurface air temperature only aboutone-halfthedifference is at the equator during equinoxconditions.As onewould expect,the difference betweenthe surfacetemperature and the air temperature greatestin low- and is mid-latitude southern hemisphereummer henthe amountof solarradiationreachs w ingthesurface greatest. is The maximum andminimumdaily temperaturesswell asthe number hours a of per day above273K for severalatitudesand seasons shownin table4. From l is thesedataonecanestimate that duringtheMartiansummer, ndprobablyfor about a one-halfthe fall and springseasons, predictedtemperatures the would be above freezing about25%of thetime. Despitecertainambiguities,he Mariner t experiments aveprovideda wealthof h data on the temperature structureof the Martianatmosphere. They alsosupply importantdatato checknumericalmodels. heradiooccultationexperiments ield T y atmospheric temperatures nearthe surfacebut do not giveactualsurfacetemperatures.However,hesecanbe determined t from radiometerresults. orexample,he F t near-surface temperature determined from the Mariner6 entry into occultationwas 250K at 15h45 Martianlocal time at 3.7 N. This measurement m corresponds to northern hemisphere Martianfall, about one Earth-monthafter equinoxpassage. The radiometerreporteda surfacetemperature 269to 279K; a discontinuityof of TABLE DIURNAL 4.TEMPERATURE EXTREMES
ABOVE FREEZING FOR SELECTED [Temperatures Location (Northern Hemisphere) Equator Polar (75 ) Mid-latitude Mid-latitude Source of data: Season in parentheses ON MARS AND HOURS LATITUDES AND SEASONS PER DAY

refer to a snow or frost covered surface] Maximum surface temperature, K Minimum surface temperature, 155 130 (145) 170 140(145) K Hours above freezing

Equinox Equinox Summer Winter

295 205 (175) 290 210(183) (1968). 18

Gierasch and Goody

about 25 K, which is reasonably closeto the theoreticalvalueof 35 K estimated from the study of Gierasch andGoody.Their calculated groundtemperature corresponding 15h45 localtime is 270K, very close to the radiometer to m results. During
the exit of Mariner at latitude not show 6 from occultation, at the temperature Mariners surface of a near m local during about 6 and surface time. this 150 7 agree air temperature A large temperature time K. and, indeed, the the of 164 Gierasch and K was and nearstudies reported ity would Goody surface 79.3 N at 22h10 discontinu-

be expected a surface

Thus, well with

surface modeling

temperature

data from

of" Gierasch The the Martian

and Goody. Mariner year, the surface 30 9 mission from planet data K lower of etal., their early was during than the provided northern obscured northern predicted dust, temperature hemisphere by dust data over a significant the part data summer of the in the portion of

winter during

into

season. mission For 10gin model.

Unfortunately, so that example, region After Mariner Goody the is some apparent but data is difficult that study

the early winter radiometer

temperature

hemisphere from the from late overall given

may 6 and

be atypical. 7 thermal

temperature

as deduced

Mariner winter, than with the the

clearing were (Kieffer since the and in still

during lower are not

surface model

temperatures from the early and of Gierasch

increased,

somewhat 1973). results 9 data than and

predicted winter those the

A comparison

for a late lower

season. predicted previous

However, in the Mariner (Kieffer

it is obvious modeling

Mariner also mid-

are somewhat those

than during

lower

obtained in the

missions, especially et al., 1973). SURFACE ENERGY

high-latitudes

southern

hemisphere

BUDGETS

As missions, planet ing the by poles. by

a result

of

the

low

Martian of the footnote

surface possibility 1, page of

temperature of increasing energy 63) infrared transfer surface current the fluxes. might

reported

from

Mariner on the

it is natural changing the suggestions can be the greenhouse

to think surface (see

temperatures For example, surface to from balance the

and atmospheric emission the the nature alter the

increasground. to such be

effect

raise radiation

temperature

enhancing proposal Both

atmosphere's suggests evaluated augmenting would

Another proposals understood.

advective of

of heat energy

equator Before must

planet's

energy

balance.

Physically, system inventory energy heat to the

temperature net energy inflows Martian convection

may absorbed and

be

viewed by (and

as net

the

thermal done an energy

response on) the budget. 3. During flowing

of

a given An

work

system. A schematic the daytime,

of all energy budget for the and

outflows

is called

surface (G,

is illustrated tt, and 19 E) are

in figure usually

conduction

away

from

the

ATMOSPHERE

'r

SURFACE

SUBSURFACE La L I G

Figure 3.- Schematic energy budget for absorbed by the surface; L a = infrared the surface; L o = infrared (longwave) duction; +-H = sensible heat convection;

Martian surface; F= solar radiation from sun and sky (longwave) radiation from the atmosphere absorbed by radiation lost by the surface; +-G = subsurface heat con+E = latent heat convection.

surface (-). canic

(+).

At are such and by

night other

they minor

usually

flow

toward of the and the

the

surface,

but budget

with

less intensity have been in vol-

There action

components heat (if All from any), these

energy lost sources at

which heat surface

neglected,

as geothermal marsquakes winds.

planet's heat

interior, the are

released

in frictional to be small

dissipation enough servation lost by the

surface

energy

believed

to be neglected. Assuming that the mean surface that energy temperature absorbed remains by the constant surface for a year, is equal conof energy surface. requires Hence, to energy

F + L aTo used: make use of (1) the terms must

L o = +G +H +E be estimated. The following relationships

(1) are

F = (1 -A) where surface Mars A = surface after having albedo, been in table and Q = solar attenuated 3(e). by

Q (direct + diffuse) values incident

(2) on the and

radiation the

atmosphere:

of Q for

Earth

are provided

L o = eaTo4

(3)

2O

wheree
(=1.357

= surface 10 -12

emissivity ly/sec/K

(usually 4 =5.6710-5

0.9-1.0);

Stefan-Boltzmann 4) (1 Langley, the

constant ly= 1 cal/

erg/cm:/sec/K (K). This

cm 2); and T o = surface temperature radiation law for a gray body.

is simply

Stefan-Boltzmann

L a _ KL, where K is an empirically well near on the Earth derived for constant. averages. water But even This relation with a value the of 0.73

(4) holds of this does 12%.

remarkably value. Only

annual where 0.73.

Table

5 shows

derivation significant, error is only

equator, from data

vapor there

emission the

becomes

K deviate Table tations priate greenhouse

significantly similar

percentage based of 0.12 of on

6 provides for for a pure K for effect.

for the

Martian

atmosphere A value

radiation seems

compuapproof the

carbon current

dioxide Martian

atmosphere. conditions;

to 0.15 the

K is a measure

strength

G=0,
over heat an annual over energy from period, and one and large region approximately water bodies to another. zero where However, over this a diurnal does not period. apply This may

(5)
not

be true

oceans

currents

can transport

considerable on Mars.

TABLE

5.-

ESTIMATION

OF K FOR EARTH, RADIATION FLUXES L a, klya/yr 117 135 145 155 175 185 210 245 250

BASED

ON OBSERVED

Latitude, deg (Northern Hemisphere) 80-90 70-80 60-70 50-60 40-50 30-40 20-30 10-20 0-10

L o, kly/yr 160 190 2O5 220 250 260 285 310 300

K, = La/L o 0.73 .71 .71 .70 .70 .71 .74 .79 .83 0.733 mean

al langley, ly = 1 cal/cm 2 Source of data: Sellers (1965), figure 14, page 42.

21

TABLE

6.-

ESTIMATION

OF K FOR RADIATION

MARS, BASED MODEL L o, ly/day 279 283 286 288 290 294 303 307 309 developed

ON THEORETICAL

CO2 partial pressure, mbar 3.75 5.1 9.0 12.8 16.7 36.0 159 278 397 Source of data: Michigan). Computer

L a, ly/day 33.8 43.3 48.2 51.9 54.8 61.2 79.4 86.0 90.8 model

K, = La/L o 0.12 .15 .17 .18 .19 .21 .26 .28 .29 by W. Kuhn (Univ. of

H and E

are

difficult

to

evaluate

since

both eddy

require transfer possibly then

knowledge coefficients, in polar

of

fine-scale neither

atmospheric is well all other

turbulence determined for terms Mars on

structure, in a simple an annual

or of bulk way. basis.

of which since

Except H can can the now energy at

regions,

it is safe

to set E = 0

be estimated with annual for Earth. First, values role that on

as a residual, the above energy

in the As an

energy application for current

budget of

be estimated budget, three data features.

parameterizations. fluxes The on Mars are are pre-

calculated sented assumed equator. except

conditions with reaches convection it reaches This the

different

latitudes.

results

in table Table 7 zero Second, at the

7 together reveals on some Mars, heat

comparable appreciable plays -36

interesting

subsurface Earth, energy

heat particularly balance

flux,

while at the

a small ly/day, value,

in the

of Mars, from value

pole

where

is, transferred even the

downward Earth's to large polar

atmosphere by a factor and

to surface. of 1.3. may Being

is a large residual

exceeding however, Third,

term,

tl is subject the

cumulative

errors,

only

be qualitatively

correct.

magnitude

of atmospheric

radiation absorbed and is an indication the latent heat although flux

by the surface (L a) is 17 to 26 times larger on Earth than Mars, of the vastly more powerful greenhouse effect of Earth. Finally, (E) term assumed contributes zero for Mars. significantly at lower and middle terrestrial

latitudes,

22

TABLE 7.-

ANNUAL

SURFACE Equator, ly/day

ENERGY

BUDGETS 45 N, ly/day

FOR EARTH

AND MARS

Heat flux

North Pole, ly/day Earth 93 320 438 -6 -27 8 Mars 40 12 88 0 -36 0

Earth F La Lo G H E 427 682 821 60 30 198

Mars 248 39 281 0 6 0 Sellers (1965),

Earth 337 479 685 -19 46 104

Mars 168 27 192 0 3 0

Sources of data for Earth:

table 16, page 103; figure 14, page 42. 0 and 45); A = 0.75 and North Pole,

Constants used in Mars energy budgets: A = 0.25 (for latitude for North Pole; e = 1.0; T O = 220 K, 200 K, 165 K for 0,45, respectively;K = 0.14 for all latitudes.

This tions with

energy time at the Lettau

balance can be

approach explored of The

provides by the

estimates method and of

of average "climatonomy" in

fluxes.

Flux

variaby by the variaof as

developed form

H. Lettau Lettau and The tions time, Fourier constants with F(t). quantitative

University (1969).

Wisconsin term

described was

preliminary

"climatonomy" problems.

coined surface surface

to emphasize temperature as a function budget By not

approach basic time, The goal

to climatological of thermal given the

climatonomy absorbed To(t) can and be

is to predict radiation the heat at the

To(t), method series these

expresses with constants elaboration

fluxes and and is lengthy

in the energy angles. and does related to the

cosine

unspecified of the

amplitudes determined technique the

phase

suitable unknown warrant paper by of

parameterizations

of To(t ) . Full

a detailed account here. (The C. R. Stearns of the University The the A energy following fluxes variables in the most

theory will be of Wisconsin.) need general

subject for

of a forthcoming

to be specified case:

complete

parameterization

= surface

albedo

= Bowen

ratio

= H/E,

if E 4:0

Cp F

= specific = mean

heat solar

of air radiation absorbed on the surface for a given heating cycle

23

AF K n To

= amplitude = La/L o = heating

of F

cycle frequency

(e.g., annual

or diurnal)

= mean surface temperature over the heating cycle (this, however, can be generated by an iterated process starting with an initial guess of T o ) = surface = surface friction roughness velocity length cycle

u* zo 6 e

= phase angle of the heating = surface = surface emissivity thermal

admittance

= (XC) 1/z

(X = molecular

heat

conductivity,

and C = volumetric p = air density The tures based on climatonomic a machine

heat capacity

of the submedium)

approach computation

has been

used to derive were obtained. without in which

annual

surface

tempera1) was by II) by

and ranges

for Mars. Two solutions

The first (solution atmosphere,

for a planet of Wisconsin,

performed

C. R. Stearns

at the University

heat fluxes L a, H, and E were

all set equal to zero, and To was generated iteratively. The second (solution consisted of a first-order hand calculation carried out at Ames Research Center S. Berman. Three selected latitudes in the northern hemisphere were analyzed

using

observed values of To taken from Mariner flybys and summarized in table 1tg_. Also. a dry atmosphere was assumed, so that L a and H were not zero quantities. In addition, the following variables were specified: # = 0.014 ly/deg/sec _/2 for sand surface (assumed equatorward of 75N and snow surface (assumed poleward of 75N 75 S) and 0.007 ly/deg/sec and 75 S), O= 1.26XlCFS _/2 for g/cma,

Cp = 0.822 erg/g/K (for pure CO2), u* = 250 cm/sec, and zo = 1 cm. Both solutions are summarized in table 8 below. A comparison solutions strong shows radiative good losses agreement at the at low and middle poles must be offset latitudes, considering strong of the assumptions, but poor agreement at high latitudes.

of the two the nature since atmospheric

This is not surprising

by equally

24

TABLE

8.-

ANNUAL

SURFACE TEMPERATURES FROM CLIMATONOMY Solution I - (Stearns' program) (No atmosphere) Annu_ range, K 149 107 111 60 30 11 22
4]

AND RANGES

FOR MARS

Latitude, deg 90 N 75N 60 N 45 N 30N 15N

Surface albedo 0.75 .75 .20 .20 .20 .20 .2O

To, K 103 119 184 206 219 226 228 226 220 208 188 123 108

Solution II - (Berman) (Dry atmosphere) To, K Annualrange, K 165 ...... ...... 198 ...... ...... 220
......

44

45

19

15S 30 S 45 S 60 S 75 S 90 S

.20 .20 .20 .20 .75 .75

64 93 141 152 186

...... 214 ...... ...... 173

---

---

fluxes more sphere.

(advection) realistic First-order

to

the polar

surface. regions.

Consequently, Solution were also similar at the time

solution be

II results correct for

appear

to be

in the diurnal

I would

without three No

an atmolatitudes solution in 1

variations

hand-calculated to solution of writing.

the

southern variation in table minus

hemisphere results 9. The the than number lower are daily

in a manner available temperature Thus, of 324 during for from hours threshold at

II above. The data

diurnal below mum

are displayed as the maxi-

range 45S

in table on the

9 is interpreted day of the

minimum. a high of reported

summer value may and

solstice, is about be erroexceeds estimated extrapoequinox the 1938 hr

climatonomy 20 K higher neous. 270 The

predicts

K and the

a low of 207 day that of the

K. The surface life, numbers

upper

Mariner

occultation

experiments,

temperature has been

K, a rough

survival

terrestrial these crude

assuming lated value over

a sinusoidal a full for yields

temperature year time by and hours

wave. making each

Finally, the

have been that for the

Martian half 2164 the

assumption applies

applies

solstice

value

one-fourth and

time. This at 45 S. In apmual

annual

in excess

of 270

K at the equator predicts

summary, temperature

surface from

temperature summer to

climatonomy winter of about

a difference K in middle

in mean and polar

45

25

TABLE 9.- MARTIAN DIURNAL SURFACE FROM CLIMATONOMY, ASSUMING

TEMPERATURE VARIATIONS A DRY ATMOSPHERE Daily temperature range, K Hours above 270 K

Latitude, deg

Surface albedo

Observed

To, K
Case 1: Equmox

0 45S 90 S

0.2 .2 .75

220 214 173

117 81 0 ;ummersolstice 125 117 0 wmtersolstice 97 20 0

4.2 0 0

Case 2: Southemhemisphere 0 45 S 90 S .2 .2 .75 Case 3: Southern 0 45 S 90 S .2 .2 .75 240 265 200 hemisphere 200 162 145

8.2 11.3 0

0 0 0

latitudes, slightly except diurnal above when major and the

and over during range 270 these point

about 100

half the

this

number

at

the

equator. and about

The the

diurnal same for

range mid

would latitudes,

be

K at

equator when About from the

all year, the 2000 range equator those

winter is always

solstice zero. region are

would

drop

to 20 K. At year were is general Goody

the poles

the to be

hr per Martian

estimated

K in the results

to 45 S. There of Gierasch and

agreement (fig. 2). The 45 S

compared is the

with

of disagreement number RADIATION

predicted above

summer freezing.

temperature

at latitude

corresponding

of hours

ULTRAVIOLET

Electromagnetic ultraviolet organisms. larly let Life light in the generally to cells on Earth (_ < 2000A) Nucleic region produces

radiation portion and

in the of the proteins

ultraviolet spectrum strongly

(2000

< ?_ <

3000A)

and vacuum threat to living particuultraviodamaging Lett, 1967). ultraviolet in

constitutes absorb

a severe

acids

ultraviolet which from

radiation, transfer and the solar

of 2500-2900A energetic with by

(Giese, secondary molecular atmospheric by the

1964). photo

Radiation electrons

in the vacuum

energy

by collisions is protected

constituents attenuation Martian

(Alexander

incident upon the planet. Attenuation of this radiation

atmosphere

can be expressed

the form 26

= s(X)Fg(r)T(x)
Here Is(k) atmosphere geometrical distance and the coordinate is the spectral distribution of solar radiation incident upon

(6)
the Martian 1.52RE); Fg(P)isa P (the instantaneous of the site 0, , in a Martian (_<1). Both

at mean distance from the Sun (RM = 1.52AU= factor (_<1) which is determined by the variables the Sun R, the instantaneous instantaneous latitude and latitude longitude, Sun's

from

and longitude

or, _, measured coefficient

system),

and T(X) is the atmospheric

transmission

T(k) and Fg(F) are dimensionless, and Is(k) is measured in units of ergs/cm 2/sec/3,, or in terms of ergs/cm 2/sec in a bandpass of a certain number of 3,ngstroms. A 503` bandpass is used in the analysis following sections, when considered the spectral appropriate. function Is(h), the geometrical factor It is concluded that 1974) in the Martian the measured mean too small to screen ultraviolet radiaIn the

Fg(F), and the transmission coefficient T(k), are described. while there is sufficient carbon dioxide ('80/am atm) (Barth, atmosphere to screen vacuum amount of ozone (5/am atm) effectively the ultraviolet tion constitutes a serious Spectral Distribution ultraviolet radiation effectively, is about one order of magnitude

portion of the spectrum. Consequently, threat to terrestrial life forms.

of the Martian Solar Constant with a spectral distribution similar to that of a black ultraviolet portion body at

The sun radiates about 6000

K. However,

in the ultraviolet

and vacuum

of the

spectrum there are deviations from this behavior, and more precise data are required. The spectral distribution of solar intensity incident upon the Earth's atmosphere at mean distance from the Sun has been determined These density values must be multiplied by: 1 \RM/ to be appropriate These incident data upon the for mean Martian have been Martian = distance 2 =0.433 from the Sun. the mean values of solar intensities of waveas a function (7) (Berkner and Marshall, 1965).

used in constructing atmosphere

in a 503` bandpass

length (tables 10 and 11). In the vacuum ultraviolet portion of the spectrum the spectral distribution is not well approximated by a blackbody curve. Estimates for the vacuum ultraviolet and ultraviolet solar constants for Mars are obtained from tables 10 and 11 in the forms

SvU V _ 80 ergs/cm:/sec, SUV _ 7 103 ergs/cm 2/sec, 27

( 10003, (20003`

< k < 20003`) < k < 30003`)

TABLE

10.-

SOLAR

VACUUM

ULTRAVIOLET I

SPECTRAL

DISTRIBUTION

XCAI i_(x)
700 75o 800 850 900 950 1000 1050 1100 1150 1200 1250 1300 I 0.024 I .024 i .026 I .048 I .11 1 .065 [ .078 I .043 I .026 I .035 I 2.47 I .065 I .078

x(A)
1350 1400 1450 1500 1550 1600 1650 1700 1750 1800 1850 1900 1950

i(X)
0.11 .11 .22 .41 .74 1.40 2.17 3.55 5.20 8.23 12.1 17.8 23.8

aValues of ICA)shown give the number of ergs/cm z/sec in a 50)_ bandpass centered at the indicated wavelength incident upon the Martian atmosphere at mean distance (1.52 AU) from the Sun. TABLE 11.SOLAR ULTRAVIOLET SPECTRAL DISTRIBUTION a

x(A)
2000 2050 2100 2150 2200 2250 2300 2350 2400 2450

T(x)
30.3 39.0 62.8 104 134 152 i156 I 139 147 ]169

x(k)
2500 2550 2600 2650 2700 2750 2800 2850 2900 2950

f_
165 242 303 390 450 480 650 870 1080 1300

aSee footnote Since the total integrated Martian solar

O) of table 8. constant is

Stota only about 0.01% of the Sun's

! -_ 6XI0 energy to

s ergs/cm

2/scc ultraviolet relatively the intensity and large about spectral 1.0% in the at

is in vacuum note the that

ultraviolet region. It is of considerable about 1200A in table

interest

feature increases

10, and

to recognize

of radiation

28

rapidly with increasing table 1 1. Geometrical

wavelength

in the near ultraviolet

spectrum,

as indicated

in

Considerations factor bg(P) appearing in equation (6) can be written in the

The geometrical form

t:g([') where (RM/RM) with v the true eccentricity, site latitude Maximum and solar anomaly g2 is the

= (RM/RM)2

cos _

(8)

= ( 1 + e cos v)/( 1 - e 2 ) 360), e = 0.093 is the Martian angular direction

(9)
orbital

of the Sun (0

angle between (0,4)).

the Sun's

(oc/3) and the

and longitude

and minimum (RM/RM)2max (RM/RM)2min

values of (RM/RM)2 = 1.22, u=0 = 0.84,

are obtained winter)

in the forms

(Northern

v= 180 (Northern about the mean

summer) are obtained significant as a consein the of haronly.

Evidently,

intensity

variations eccentricity

of 20%

quence of the orbital Mars-Sun distance. The monic factor factor and site latitude

of Mars which

produces

variation

cos g2 varies from zero to one, and can be expressed longitude theorem. and the Sun's angular position of latitude It is perhaps more convenient to average

in terms

using the spherical and true anomaly

addition

the geometrical

over a Martian

day, so that it is a function factor yearly

This average geometrical ----0.3, whereas significant

Fg(O,v) at the equator is approximately constant at variation is observed with increasing latitude in both about 40 that the geometrical of about 50 in the northern to reduction in mean solar

hemispheres. It is only for latitudes greater than factor is less than approximately 0.1. A latitude hemisphere intensity. Atmospheric is particularly favorable with respect

Transmission function can be written in the form

The transmission

// T(X) = I1 T i (X) i=1 29

(10)

where
Tt.(_t) = exp(-oiNi) and is the transmission absorption (cm -2 ). cross section function (cm 2), for the ith molecular and N i is the column (11) species, o(_) is the molecular density of the ith species

The absorption coefficient ), > 2200A, and the 03 absorption been determined sections using (Berkner

kiOt)(cm -_) for 02, H20, 03, and CO2, for coefficient in the 1800A to 3000A interval have 1965). These data can be converted to cross

and Marshall,

oi(X) = ki(X)/N o where N O = 2.69X 1019/cm3 (Loschmidt's number). The cross section o(_) tbr molecular ozone has been and additional available data (Griggs, 1968), and is tabulated were ozone obtain used in constructing transmission functions for

(12)

constructed from these in table 12. These data 100/am atm of to

1, 10, and

(tables 13 and 14). Evidently, at least 100/am atm of ozone are needed significant absorption of vacuum ultraviolet and ultraviolet radiation.
TABLE 12.MOLECULAR OZONE PHOTOABSORPTION CROSS SECTION a

X(A)
1000 1050 1100 1150 1200 1250 1300 1350 1400 1450 1500 1550 1600 1650 1700 1750 1800 1850 1900 1950

o(?,)
0.93 1.10 1.30 .75 1.12 .74 1.50 1.50 .74 .60 .45 .30 .15 .12 .10 .10 .082 .065 .048 .038

?,fA)
2000 2050 2100 2150 2200 2250 2300 2350 2400 2450 2500 2550 2600 2650 2700 2750 2800 2850 2900 2950

o(x)
0.030 .04 1 .052 .12 .19 .29 .42 .59 .75 .97 1.02 1.10 .97 .94 .76 .57 .37 .22 .13 .08

a6(h) x 1017

cm 2

3O

TABLE

13.-

VACUUM ULTRAVIOLET MOLECULAR TRANSMISSION FUNCTION 1/am atm 0.98 .97 .97 .98 .97 .98 .96 .96 .98 .98 .49 .99 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00 14.ULTRAVIOLET TRANSMISSION 10/am atm 0.78 .74 .71 .82 .74 .82 .67 .67 .82 .85 .89 .92 .96 .97 .97 .97 .98 .98 .99 .99 100/am atm 0.082 .052 .030 .13 .049 .14 .018 .018 .14 .20 .30 .45 .67 .72 .76 .76 .80 .84 .88 .90 MOLECULAR FUNCTION

OZONE

X(A) 1000 1050 1100 1150 1200 1250 1300 1350 1400 1450 1500 1550 1600 1650 1700 1750 1800 1850 1900 1950 TABLE

OZONE

x(A)
2000 2050 2100 2150 2200 2250 2300 2350 2400 2450 2500 2550 2600 2650 2700 2750 2800 2850 2900 2950

1 /am atm 1.00 1.00 1.00 1.00 .99 .99 .99 .98 .98 .97 .97 .97 .97 .98 .98 .98 .99 .99 1.00 1.00

10/am arm 0.99 .99 .99 .97 .95 .92 .89 .85 .82 .77 .76 .74 .77 .78 .82 .86 .91 .94 .97 .98

1O0/am atm 0.92 .90 .87 .72 .60 .46 .32 .2t .13 .074 .064 .052 .074 .080 .13 .22 .37 .55 .70 .81

31

Molecular ultraviolet photoabsorption absorption are cross 80 rn atm portion

carbon

dioxide spectrum at about

is completely (Banks 2150A. Martian that and In

transparent Kockerts, table 15 the

in 1973);

the

visible

and

near for

of the occurs section

the threshhold dioxide Since of Table

carbon wavelengths. the data

photothere 15 and function

is tabulated dioxide in the

for vacuum

ultraviolet atmosphere, dioxide

of carbon spectrum

the absorption satisfies

of CO 2 show

the carbon

transmission

TCO 2 (X) = 0 TCO2(X) = 0.1

_, _< 1900A X_ 2000A

TCOZ (X) = 1.0 Consequently, ultraviolet, Upper atmosphere These that species of ozone, but the limits have carbon on been the dioxide amounts in the of (Owen Martian radiation. various and

X _> 2000A

atmosphere minor Sagan,

is opaque

to vacuum Martian 1972). than

is transparent

to ultraviolet

constituents 1972, Pimental

in the et al.,

determined have

generally although

ultraviolet search

absorption

cross

sections

much

smaller

a detailed

is recommended.

TABLE

15.-

MOLECULAR

CARBON DIOXIDE CROSS SECTION a

PHOTOABSORPTION

X(A)
1000 1050 1100 1150 1200 1250 1300 1350 1400 1450

o(X)
0.75 .74 .74 .74 .0074 .010 .048 .093 .052 .060

X(A)
1500 1550 1600 1650 1700 1750 1800 1850 1900 1950

o(x)
0.050 .075 .069 .036 .0022 .00093

lO_ lO_
10-s 10-6

ao(x)X 1017 cm z

32

Ultraviolet

Intensity

Estimates from the region on

for the Martian previous due the data that surface, of

Surface the Martian the _-0.3 data atmosphere of tables the the is opaque To estimate I 1 and data of table ultraviolet The action in the 16. by

It is evident the been 1000A of intensity

to 2000A ultraviolet with

to carbon Martian factor

dioxide

photoabsorption. to obtain

14 have

combined 100/am

a geometrical of ozone

Evidently, approximately ation

atm order

is sufficient in the to 30003,

to attenuate 2400A

solar interval.

one

of magnitude 2700A

to 2700A and

attenuspectra

is significantly processes total The

less in the

interval,

detailed in estimating

for biological damage. 50 ergs/cm presence Martian winter observed is clear of The

of interest

will have in

to be employed the 2400A intensity as the (Barth of the to

ultraviolet is about in the of

integrated total

intensity ultraviolet

2700A

interval

2/see.

radiation Inasmuch 60_matm

is 103 ergs/cm=/see, measured 1973), that

100 #matm

of ozone.

maximum etal., fact

amount

ozone season at

is approximately the polar cap,

observed

in the damage 2/see,it polar is

and

in

light

significant 104/ergs/cm in the

in terrestrial that useful 16.-

organisms radiation

for an ultraviolet protection is presently

flux

>103 afforded

Martian

TABLE

ULTRAVIOLET 1 vm atm 03 9.10 11.7 18.8 31.2 39.8 45.1 46.3 40.9 43.2 49.2 48.0 70.4 88.2 115 132 141 193 258 324 390

INTENSITY 10/Jm atm 03 9.00 11.6 18.7 30.3 38.2 42.0 41.7 35.4 36.2 39.0 37.6 53.7 70.0 91.3 114 124 178 245 314 382

AT THE MARTIAN 100 vm atm 03 8.37 10.5 16.3 22.5 24.1 20.9 15.0 8.76 5.73 3.75 3.17 3.78 6.73 9.36 17.6 31.7 72.2 144 227 316 wavelength.

SURFACE

x(A)
2O00 2050 2100 2150 2200 2250 2300 2350 2400 2450 2500 2550 2600 2650 2700 2750 2800 2850 2900 2950 aValues

of ergs/cm _/sec in 50A band pass at indicated

33

regions.

However,

a maximum regions study present

value of 5 ttm atm of ozone the radiation ultraviolet shielding of the absorption

is more spectra

appropriate of trace

for atmostudy

lower latitudes, A more spheric dance species

in which detailed possibly

provided

is negligible.

on Mars is recommended, atmosphere in order

as is a more careful estimates

of the chemistry

of the Martian

to provide

of abun-

of trace compounds.

34

5.

THE

TRANSPLANTATION ECOLOGY a most forms optimistic present would

OF

TERRESTRIAL

ORGANISMS

TO

MARS

A MARTIAN Even able of even have

appraisal Martian be quite to determine

suggests environment restricted the

that

the

kinds and

of terrestrial limited, extent (table

organisms growth 17). There to simulated of the feel

to survive these

in the

are quite in vigor

and the

been

many

attempts

response have

of microorganisms in part conditions,

Martian

environments. choice

While

conclusions and

varied,

a reflection investigators

experimenters'

of organisms

environmental

TABLE

17.-

COMPARISON OF LIMITS ENVIRONMENTAL Range compatible Growth: 255K-377K

TO LIFE WITH SELECTED PARAMETERS with life

MARTIAN

Parameter Temperature Ultraviolet irradiation

Mars 140 K-300 K

Survival: <79K-377K

3 10 s ergs/cm 2 0-100% 10%- 100% (w/w) 0%-35% 0-13 NaC1

7 10 a ergs/cm2/sec 20 Present, amount

?

Oxygen (pO2) Liquid H20 Salinity pH Mechanical abrasion

unknown

? Dust in storms _ 10-50#m in size range of many microorganisms Not yet found Not yet found Not yet found CO, CO2, 5 mbar H20 H20

Abrasive particles >/cell size will cause mechanical damage

Nitrogen Sulfur Phosphorus Carbon Oxygen Hydrogen

Absolute

requirement

35

that there is a possibilityfor growth on Marsof certainanaerobic, cold-adapted, terrestrialbacteria. For the growth of higherterrestriallife forms,an oxygen-containing environmentis mandatory.The needfor anoxygen atmosphereerives d fromtwo functions of oxygen:it is requiredfor the formationof ozone,the ultravioletabsorbing shield under whichlife exists,andfor the metabolism higherorganisms hich is absoof w lutely dependent pon oxygen.Therearein principletwo waysby whichgaseous u oxygencanbe produced:chemically,rom wateror otheroxygen-containing f compounds, r biologically,from water.It is generally o believed the present ass that m of terrestrialatmospheric xygenwasproduced o biologically,andthat the bulk of the oxygenin the ancientprimitive atmosphere wasproduced greenplant or algal by photosynthesissa latereventafterlife hadew)lved. husphotosynthesis believed a T is to havethe capabilityof generating oxygenin amountsnecessaryo makeMars t habitable higherterrestrial rganisms. for o Photosynthesisanbe considered the fixing of carbon dioxide and water c as
into a carbohydrate organisms. is produced. dioxide, major lack water) known The is the is: In (starch) As in the presence of the of visible light and living, chlorophyllderived and from containing water, (carbon a by-product all photosynthesis necessary physical are available of higher levels be seeded of oxygen. on Mars, oxygen, (light) terrestrial

principle, elements barrier

chemical

for photosynthesis to the establishment usable

on Mars. life on Mars question grow and A critical survive,

of an atmosphere

containing

"can

terrestrial oxygen?" attempt are:

photosynthetic

organisms

generate Any Among on these desirable Over Can how the What Mars'?

to establish would

terrestrial be required which they have

organisms

on

Mars

raises

many

questions. to grow possess about generation? grow? of of interval

them Do

characteristics any

of organisms occur the

in order

for them

microorganisms If so, in the on do Martian the

in terrestrial metabolic for example, could to useful permit

environments to bring oxygen

characteristics? alterations large Martian

capabilities

environment, surface

an area

Martian be altered present humans

microorganisms the introduction What

environment if the before a number

in order

desirable time might As growth The obtained growing are grow

microorganisms be required earlier

environment could inhabit

is too hostile? the planet'? performed has bacteria

noted of

of experiments Martian was indicate if sufficient whether conditions, 36

have

been

in which been are attempted. using capable Such

the

microorganisms recent from study Mariner Martian 9. of

in simulated this His type results

environments by certain Foster that

most

performed

(1973)

data of

in the

environment as they

nutrients or not

are provided. |errestrial in order forms

studies can some

valuable under

inasmuch a particular

indicate

of life

set of extreme

ltowever,

to obtain

indication as to how terrestriallife might fareon Marsit is alsodesirable study to natural terrestrialenvironments hichsimulatethe Martianenvironment sclosely w a aspossible. he terrestrialenvironnlents T which appear ostnearlyto resemble m the general Martianenvironmentirethe cold,dry valleys f of'Antarctica.These valleys, n a areaof severalhousandsquare t kilometers, avea meanannualair temperature h of 248 K to 253 K. In summerthe air temperature doesnot rise much higherthan 273 K: however_ groundsurfacecan reach288K or higherfor a short period the during the day. Liquid water is scarceand is the principallimiting factor for the growthof microorganisms. Therelative humidity canbelessthan45_ andthe water activity of the soil0.45or less. Thereis a concentration microorganisms of aroundsources f wateranda rapid o thinning out in the locales highestaridity {Horowitz et al., 1972). According of to
Ugolini frost, beneath meter. the (1970) provided the The the most reliable the source maximum of soils the not of soil depth ranging moisture of thawing. from a few by snow where ice loss is the This ice-cemented hard permafrost to more than on the depth ice-cemented in summer to a high to the rate permalies a of layer and of atmoshown which are related the amount are to it is within ground moisture permafrost 15 cm the and the below soils. valleys the is even of area on terrestrial of the Mars of Antarctica environment hostile. organisms, planet's appear diurnal dissemination ice fields essential encumber Although certain The are extremely in many Mars types major severe respects. presents might obstacles high be environments The Martian of which appear of the overlaying eventually permafrost ions freezing. in the surface content below soil.

at depths

centimeters depends the occurs due

covered

topmost

soil surface. some the recede upward melting water

In areas of the moves

is within moistens evaporation sphere that the

surface> enters to move The

Liquid

upwards This surface.

atmosphere. from the

of water

causes sodium

Ugolini

{1970)has

chloride of moisture

in the soils,

soil and which

even

at temperatures factors deterlnines

continuously amount of life in the The to resemble dry

climatologic

pedologic in turn

Antarctic

Martian more

environment, to over the a growth

however, growth limited organisms and might lack pounds planet. The water. extant of rapid, also

serious capable

constraints

surface. to be

to the growth flux Factors on the of ultraviolet as yet Mars to be presence

of terrestrial radiation,

paucity

of water, variations.

extreme make of

temperature of close to terrestrial enough

undetermined The a the comon

organisms to the lit'c, or

unsuccessful. easily of organisnis melted, toxic

subsurface would further

surface

deficiency

elements

terrestrial

attempts

to establish

terrestrial

single

most the

important with

limiting respect

factor to water

appears

to be

the

lack

of available to that

At best in the

situation dry

on Mars may mentioned,

be analogous certain

Antarctic

valleys.

As previously 37

observations

have

been

interpreted

as indicating

the existence

of subsurface

water

ice fields

or

permafrost Antarctica. planet,

on Mars (Fanale and Cannon, 1974), as is the case for the dry valleys of To cause melting of enough ice to moisten the soil over large areas of the warming will have to be achieved. on the depth The amount of warming needed ice depends of the frozen water. on Mars would be algae) seeded on amount of the of shielding organisms

a general

to melt the subsurface As previously lethal to exposed Mars will need ozone is produced

discussed, terrestrial from the

the flux of ultraviolet light incident organisms. Cyanophytes (blue-green until a sufficient generate. Protection oxygen they

protection

from this radiation

against ultraviolet radiation might be afforded by various endogenous or exogenous mechanisms. The former might be accomplished by means of more efficient mechanisms of repairing or other entrapped ultraviolet by genetic damage, "overproduction" If purines of purine or pyrimidines reduction or pyrimidine were excreted bases, and absorbing pigments.

the sheaths

of the organisms,

considerable

of the -=2600A

radiation could algal populations

occur. Exogenous protection might be achieved by the growth of under thin layers of translucent soil. The overlying soil would not

only protect the cyanophytes from ultraviolet radiation, but probably also would retard desiccation. The thickness of the soil layer would be critical; sufficient visible radiation intensity the must be transmitted radiation to permit photosynthesis, reduced. but at the same time the of 2600A must be greatly

Sagan and Pollack (1974) suggest Martian surface has been reduced light is 3.8 102 ergs/cm 2/sec. from ultraviolet introduced of radiation on Mars must effective

that the ultraviolet flux about 0.8 cm below to 1.7 10-3 ergs/cm2/sec while the flux of Thus subsoil growth might provide photosynthesis. DNA repair or adequate While the mechanisms, is

visible

protection organisms

while still supporting have highly efficient either

the importance emphasized. LICHENS

shielding,

exogenous

endogenous

AS POSSIBLE

MARTIAN

BIOTA candidates for "seeding" to the survival conditions. of Mars because of any terrestrial Consequently, are exam-

Lichens they certain possess

must be examined certain attributes

as possible

which are requisite with respect

life under the currently of their

envisioned

Martian environmental

characteristics

to these extreme

requirements

ined. Lichens are, in essence, composites of fungal- (mycobiont) and algal- (phycobiont) cells, which live in a symbiotic relationship. The two types of cells growing together develop a "body" or thallus, which is not characteristic under of the independent on the basis of growth of either component separately. Terrestrial lichen genera are classed their general morphologies (Jahns, 1973).

three broad categories

38

The crustoselichensmay be considered most "primitive" or the least the differentiatedform. They grow in intimate contactwith their substrates are and often "inseparable"from them. The mycilia of the mycobiont spreadover the colonizedsurfacein a thin filamentousmat whichencloses algae. healgalcells the T generallyoccupythe upperpart of the thallus,the surfacelayersof which form a cortex,composed necrotic,gelatinizedcells.This type of lichencangrow comof pletely within the substrate as,for example, heendolithiclichenswhichgrowinside t rocks.The fungalhyphaeof theseorganismsxcrete"lichen substances" e whichcan dissolve stoneandallow the penetration the fungiandalgaeto depths several of of millimeters (Sayers andIskander,1973). An intermediate, oredifferentiatedlichenis termedfolicose.Folicose m lichens consistof thalli whichareformedby flattenedlobes.Theymaybeplatelike,andare attached a centraldiscoidholdfastor umbilicus. by The fructicosegroupis the mosthighlydifferentiatedlichen-type. hey consist T of strap-shaped threadlike or lobeswith a radialthallus.Somefructicose lichensmay degeneratet the baseandbecome a completelyfree.Theymayalsobedislodged by the wind andblownovertheground. All three types,as well asgradations betweentypes,arefound intermixedin the terrestrialenvironment.Furthermore,the propertieswhich aresubsequently discussedrecommonto mostlichensregardless their growthform. a of The mechanisms hich regulateand control growth in lichens are mostly w unknown(Jahns,1973).Growth is usuallyrestrictedto the tip of the thallusand rarelyexceeds few centimetersadialincrease year(Richardson, 973).There a r per 1 is negligible intercalarygrowth,andthe inner partsof the thallushaveno means of transportingphotosyntheticproductsto the growingouterportions(unlike higher vascular lants).Therefore, nly the photosyntheticproductsof the marginal arts p o p of the lichenthalluscanbe used the growthprocess. for Lichens areextremelytolerantto a lackof water.The minimumwatercontents which lichen thalli can sustainandstill remainviableafterrehydrationis between 2 and 9% of the dry weight.Waterseems be tightly bound to the cytoplasm to (Kappen,i 973).Theability of lichensto withstand desiccation not indicativeof a is capabilityto resistthe effectsof drasticwaterloss.Lichenstendto rapidlyachieve waterpotentialequilibriumwith their surroundings. A numberof isolatedobservationserveto illustratetheability of theseorgans ismsto toleratedrought.Several pecies s werekept dry for 1.5yearsin permanent light and thenfor another1.5yearsin permanent arkness. d After 8 daysof cultivation on agarmedium,thesespecimensroduced p aplanospores (reproductive spores) (Raoand LeBlanc,1966).Investigations indicate,however, hat therecanbea loss t of drought tolerance several pecies ue to long periods moistcultivation.The in s d of measurement photosynthesis lichensunderdesiccation of in stress hasbeenshown 39

to

be

variable

among rates have

species. with

There

is a general (Kappen, periods

tendancy Various alter

toward

depression have

of been

photosynthetic observed for ing. tion (Butin, water humidity more because (Buttner, water tion, to adapt example, Tile upon

drought full viability

1973). capacity

species Antarctic several

themselves regained

to extended photosynthetic upon seem without many

of desiccation.

lichens, of dryat lower o1" desiccalow water water (higher as the may air be

weeks

ability

to regain

rehydration to be several the active lichens

is enhanced observable actively (imbibition) water does

by storage effects with potentials respiration vapor uptake of in by Many the liquid

humidities contents:

(Bacquerel, metabolic most 1954). show

1948). processes: active

There growth

metabolize uptake at higher than

of liquid

Photosynthesis but has a steeper (Lange, for

is generally gradient For

contents)

of increase some than than able species

increases profitable respiration 1971). condensation since

1969).

water

photosynthetic seeins to be are

gain higher thus and by

is strong photosynthesis

uptake

water case

latter

Many

lichens thallus

to photosynthesize water (-115 intake. atm), species.

means lichens by salt

of deware also incrustaperiods of

in the very low

vapor

halophilic, do These desiccation: rchydration. Martian tively

osmotic

potentials that their a caveat

induced

not drastically observations they There colonization. can

reduce

tile viability

of coastal lichens

demonstrate fully regain

can be resistant

to long and terrestrial

ability to the

to photosynthesize use of present lichens their responses with

respire forms

upon for

is, however, Even potential exceeds the 75% have tolerance

most may shown

drought-tolerant to reactivate be hostile favorable

appear

to need Areas 1973).

a relawhere

high

water rarely

in order

photosynthesis. I Kappen, to extremely drying.

air moisture

for lichens

A number tares species, diately (Kappen, Lunge tile and

of lichens The

low temperathree imme-

freezing. after

appears

to increase normal remained to carbon

For example, uptake ahnost

cooling rewetting They

to 77 K, showed at 283 K, and been recorded and K been Metzner in several observed to visible

dioxide for

after

active

several

weeks

thereafter 273 K. of and

19731. (1969), for rate. dioxide at 266 response

have

shown

photosynthesize at 268 Biosynthesis et al., 1966). radiation show to high is that some

well,

below

example, Lange at 262

photosynthesis { 1965)observed species. (Godnev and

K proceeding of

at 50_/_ of fixation

maximum

photosynthetic

14 carbon /3-carotenes The

chlorophylls

K have

of lichens to

ultraviolet lichens lichens

is also favorable

an important characterradiation have a

environmental istics. results tendency also been llabitats The

parameter c',posurc of effects.

be considered, terrestrial

various

intensity the

visible

in variable to

A common color,

observation when may (defense) 4O exposed be a factor lichen

phycobionts sunlight. the observed,

"bleach," that

or lose

to full summer in determining thallus has been to strong

It has and [:or

suggested of certain

illumination Pigmentation response

endolithic

lichens.

of the

is suggested

as a possible

of lichens

illuminalion.

example, response

different

colored

thalli

were intensity

observed within

in the the

sz_me lichen The

species

in high

to a variation

in light

habitat.

observed

tolerance of several species to ultraviolet radiation is of particular interest. have been reported as having an extraordinary ability to withstand intensive let radiation. at 2.5 106 (Siegel present and molecular The continuous ergs/cm 2/rain Daly, repair in some 1968). bacteria, exposure of at least one species in no observable by ultraviolet to radiation effect radiation, upon for 24 hr resulted for damage

Lichens ultravioof 2537A

respiration Although as those may

The basis for this resistance have not been investigated,

was not explained. such it is possible

mechanisms

that lichens

derive all or most of their increased tolerance from morphological adaptations. A thickened cortex (upper layer), for example, has been observed in organisms which were exposed to intense visible and ultraviolet illumination. Additionally, (normally (usually tous hyphae formed certain species live on quartz This surface blocks and sand where forms of black, the medulla surface filamenthe middle which or central region of the lichen layer). thallus) consists the upper short,

by the cortex become

interspersed

with grains of quartz,

lime, or the like. The

phycobiont layer remains underneath which ill turn acts in place of a cortex screen out harmful wavelengths and against excessive ultraviolet radiation. terrestrial water organisms

and pressed closely to this quartz surface, layer. Such a protective device could easily provide the lichen with an excellent defense upon

In view of the fact that the lack of water on Mars, it is instructive of lichens. relation properties

may impose to summarize water

a severe limitation

some of the absorptive

As was stated,

loss is a rapid and relatively

uncontrolled other hand, uptake by

process under environmental conditions of extreme drought. On the water adsorption by a dry thallus is also rapid being similar to water a hydrophilic gel. Lichens can function by absorbing water vapor,

although a major portion of water is probably only gained in this fashion from atmospheres in which the relative humidity exceeds 90% (Blum, 1973). It has been demonstrated that air-dried thalli in a latent state can become reactivated and reach photosynthetic symbiosis rates close to optimal protection from values. its fungal It appears partner, that the algal member which provides of the to derives a buffer

rapid changes in humidity (Quispel, 1959). It is known that blue-green algal phycobionts can hold large amounts of water in their thick gelatinous sheaths. Furthermore, some lichen forms have "xeromorphous" structures such as a thick and solid cortex or cortical hairs (Blum, 1973). The final determination of viability is made by the resistance to irreversible damage of cell structures, which are associated with vital biochemical functions during the desiccation-rehydration The principal task of any organism used to colonize cycle. Mars is to enrich the

atmosphere in oxygen. It is therefore necessary to examine some of the important factors which might influence photosynthetic production in lichens, as well as to 41

elucidatesomequantitativeaspects the process. of Variousmaximumnet ratesof carbondioxidefixationhavebeennotedby differentinvestigators. Reid(1960)gave a rangefor the maximumnet photosyntheticrate as 0.34-3.2mg CO2/50cm 2 surfaceareacovered/hr. imilarly,optimal ratesfor somealpinespeciesrelistedas S a 0.30 0.38mgCOz/gmdry wt/hr (Bliss& Handley,1964). The moisturecontentmayhaveanappreciable intluence uponphotosynthesis. Lange(1969) found that net photosynthesis f desiccated o thalli increased rapidly with hydrationup to about60%saturation (i.e.,60 mgwaterper 100mgdry wt) in onespecies. hydrationcompensation oint occurredat 20%of the waterholdThe p ing capacity at 283 K and 10,000 lux (_2.5 ly/min). Adaptive responses to environmental open thetic conditions habitats which limit the process Arctic, which have been observed. Lichens isolated from to 10% and in the Canadian 1971). for example, moisture had a maximum content lichens carbon content net photosyndropped assume a state assimilation

rate at 309_ saturation, and Rouse, animation

fell slowly as the water there is no measurable

(Kershaw

Below a critical

of suspended

during which

respiration is extremely low. The amount of water required among species. Lange (1969) found

for minimum that

photosynthesis species,

varies considerably which were moistened

two lichen

nightly by dew condensation, photosynthesized for 3 hr after sunrise, and then dried, crossing the compensation point (carbon dioxide emitted for a short period). No carbon dioxide was detected for the rest of the day until the thaili became moistened again at night. The carbon dioxide balance averaged 0.54 mg CO2/g dry wt fixed over 24 hr. Lange (1970) emphasized the fact that the annual photosynthetic gain in certain lichens would allow for a thallus growth of 5-10%, which could be attributed The effect to be variable (1969) found of temperature, almost exclusively upon to dewfall. photosynthetic rate, although obvious, seems of light intensity

with species. Light saturation values tend to be relatively high. Lange that a desert lichen reached light saturation at 20,000 lux (_5 ly/min) the fact that the light compensation point of lichens is a function near the variation and the magnitude of these points at temperatures

at 275 K. Despite

273 K are small. Light compensation points have been shown to remain in the range of 200-300 lux (_0.05-0.075 ly/min) between 268 and 275 K. These effects are most probably due to an increase in the respiration features cycles rate of the which fungus at higher to temperatures. In summary, their evaluation 1. Resistance

some of the important for Martian colonization and to long periods

of lichens

are pertinent are summarized.

or planetary

engineering

of desiccation, imbibition

particularly by the thallus.

at low

temperatures (shows dormancy). 2. Relative ease of both water

loss and water

42

3. Requirement or liquid wateror an atmosphere f of

(>90%) 4. for rehydration of the desiccated thallus. Ability to retain a near maximum photosynthetic

high relative

humitidy

activity

in the hydration loss,

phase of the desiccation-rehydration cycle, and adaptation to such conditions. 5. Ability to show net photosynthesis while sustaining a major water although 7. extremes 9. rates decrease Reproduction with increased by means desiccation. at temperatures which as low as 260 K. are resistant 6. Ability to sustain net photosynthesis of spores easily. radiation. at low temperatures. prohibit photosynthesis. of lichens to survive rates.

to environmental

and are disseminated Extremely

8. Tolerance

to high levels of ultraviolet slow growth

10. Low photosynthetic 11. Ability to degrade 13. Existence A major environmental

light compensation points or weather stone substrates. under forms. dealing the ability with conditions which

12. Low rate of respiration of halophilic omission extremes

in experiments is concerning

the

adaptation organisms

to and

of these

grow under anaerobic conditions. Data on this property are most important evaluation of lichens for use in Martian atmospheric alteration programs. CYANOPHYTES The group photosynthesizers AS POSSIBLE MARTIAN BIOTA

to the

of organisms which appear to have the greatest potential as oxygenic on Mars are the cyanophytes (blue-green algae). As a group they

are versatile in their physiology and display considerable latitude in their environmental tolerances. Table 18 summarizes the observed environmental extremes for growth and survival of the cyanophytes (also see pages 77 and 78). Their ability to grow or survive in hostile environments and carry out oxygen-evolving sis suggests that this group of microorganisms should be considered dissemination reflection important ments activities on of their Mars. The wide distribution and because organisms, of cyanophytes versatile physiology. and because they ecological colonizers austere conditions tolerance for other which photosynthefor primary Earth is a are

on the

Cyanophytes

as primary are too about bring

on Earth

are able to grow in environtheir physiological of other soil to the growth

which

are conducive

organisms. These activities result in an increase of the soil. A gelatinous sheath aids in binding soil moisture the in and growth polar prevent regions. erosion. stimulate identified of vascular plants.

in the humus and combined nitrogen soil particles which helps to maintain also genera produce described substances mats of which Nostoc of cyanophytes have been

Cyanophytes Several marble 43

Holm-Hansen

(1963)

10- 15 cm in depth

on top of alkaline

areas on Ross Island,

South Victoria

TABLE

18.

OBSERVED ENVIRONMENTAL EXTREMES FOR SURVIVAL OF BLUE-GREEN ALGAE Lower limit

GROWTH

AND

Upper limit 373 K (dry soil) g 363 K (wet soil) g 346 K (neutral and alkaline hot springs) c pH 10-11 pH 13 (plectonerna Eh: +700 mv (pH 4) +670 mv on silica) h

Temperature

(survival)

4 Ke

Photot rophic metabolism

264 K (Saline Lake, U.S.S.R.) 243 K (lichens) d pH 4.0 c

Hydrogen-ion (growth) Oxidation

concentration

potential

(growth)

Eh: -200 -184

mv (pH 6) mv (aphanocapsa thermalis ) (pH range 1.2-6.6) (0.001%)

Total salinity (growth)

Freshwater

27.5% (Great Salt Lake) b 31.5% (Dead Sea) b >5.2 km above sea level Mts.)

Altitude

(growth)

0.396 km below sea level (Dead Sea) b and Schizothrix and Anacystis

(Himalaya round at Atacama

Resistance to aridity desiccation

Desert, Chile

Nostoc revived after 107 years of storage as a dried herbarium specimen e Some blue-green algae survived more than 106 fads of'),-irradiation when administered at dose rates of about 2X l0 s r hr-_ f Microcoleus survived 2560 kr 7-irradiation eCameron, fGodward, gLund, R. E. (1963) M. B. E. (1962) from Co 6 o source

Resistance to ionizing radiation

aTable modified

from Schopf(1974). unless

Data

adopted directly from Schopf otherwise indicated bBrock, T. D. (1969)

J. W. G. (1962) J. R. (1963)

hVallentyne, CBrock, T. D. (1973) djames, P. F. (1955)

1Stewart, W. D. P. and H. W. Pearson (1970)

Land. about Islands. been

Fogg 2 mm observed

(1973) below just enough

found the

that

a distinct of white abundant sand

zone

of

cyanophytes

was

often of the

present Falkland have

surface that the

quartzite and on

sands areas the

on the coasts growths of the of west

He also

reports below light are

healthy

cyanophytes coast to allow of time. algae

surface between

of Scotland. photosynIn some

Apparently thetic growth.

penetrates to survive alternate but when

sand for

particles long periods

Cyanophytes regions while appears where water arid

able

desiccation with the moist

conditions

conditions becomes

the moist,

are inactive growth which is

is unavailable, time.

soil again readily 44

extensive on lava

in a short

Cyanophytes

become

established

devoid nitrogen elemental extremely

of other being nitrogen

growing especially coupled

organisms, suited with for

those growth

species on

which such confers

are able

to fix elemental The ability latter to fix species

material.

photolithotrophy

on these

simple nutritional

requirements.

All of the cyanophytes are phototrophs which utilize carbon dioxide either exclusively or almost exclusively as their source of cellular carbon. An absolute requirement marine Provasoli, compounds, shown that ties which for organic carbon has been species demonstrated in only a small number vitamin B_ 2 (Pinter et al. (1971) are able to assimilate Van Baalen growth of and have cyanophytes, and certain barely and these all appear to require

1958; Van Baalen, organic support

1961 ). Many cyanophytes compounds

organic

in some cases to grow in the dark. photosynthetic growth

are able to stimulate

at light intensiAlthough

(1.4X 104 ergs/cm 2/sec).

cyanophytes are capable of photosynthesis oxygen concentration stimulates carbon (1970) demonstrated the conditions of anaerobiosis ance they

in the presence of oxygen, a reduction in dioxide fixation. Stewart and Pearson

photosynthetic growth of Anabaena flos-aquae under (see also Weller et al., 1975). Notwithstanding the toler-

of blue-green algae to diversified environments, it does not appear likely that could grow extensively even in the areas on Mars where the most moderate conditions These prevail. This express conclusion the view is supported that the by the observations (Horowitz of et al., a et al., on the soil biology investigators isolated from the most of the dry valleys of Antarctica viable valleys arid parts of these

climatic Horowitz 1972).

organisms,

including

cyanophytes,

do not constitute

population attuned to growth in the prevailing environment. Furthermore, they present data indicating that much of the soil in the valleys is sterile (20% of 328 soil samples were sterile as examined by plate count). It should again be emphasized than appears from also that the dry valleys represent a much less hostile even the most temperate regions of Mars. The permafrost soluble salts subsurface into the toward habitat the assumes soil. surface. overlying This environment to exist in the melting carry waterthe reduce

a movement Moistening

of liquid water of water thereby of the soil would

movement

would

amount of dust entering the air due to wind action, energy to reach the surface of the planet. BIOLOGICAL In a closed occur. inanimate net result cally form CYCLING system, OF ELEMENTS recycling

permitting

more radiant

ON MARS elements essential to life must The

of the chemical cycles circulate

On Earth, biogeochemical into organic, molecules of these

these elements equilibrium and

from the inorganic, back again. for such biologi(figs. 4 and 5).

protoplasmic

combinations a steady-state sulfur, oxygen,

and then carbon

cycles is to maintain as nitrogen,

important

45

O_

NO2

--

,, _ _9_P"

DE4/14 R NH2 _

2"/_

N2

NO3 ...... NH3

_\"

AEROBIC ANAEROBIC

_\

"4_/_'..,.,,..._

R NH 2 O r,

/O

/_

NO 2 -

N2

Figure 4.-

The biological

nitrogen

cycle.

AEROBIC ANAEROBIC

Figure 5.For example, the by the removal efflux of carbon of carbon

The biological dioxide dioxide from

sulfur cycle. the atmosphere by respiration: by photosynthesis

is balanced

produced

Photosynthesis CO2 + Hz 0 // "_ C(Hz O) + 02

Respiration

Key On

to the the

transformation most soils

of these contain

elements a large 46

from

one

form of

to another microorganisms

are microbes. (bacteria,

Earth

number

protozoa,andfungi). Their
and ments life tinually simpler organisms. if would cyanophytes cause the alone accumulation and inorganic available. l'orms, guided flow into compounds It is through by many reservoirs forms

metabolic which the of thus

activity

results for

in the modification biologically raw but for the materials are broken

of organic important elelower conto of

is essential complex unusable made

making that

interaction factors, compounds, available

of numerous

higher do

and not

environmental

down growth

inorganic

and

continued

were

to be

introduced matter

to Mars, containing be most

their carbon,

metabolic nitrogen, to prevent weight organic

activity sulfur, such comon on as a

of organic hydrogen. into

phosphorus, one-way pounds Earth Mars, flow by depends as well

oxygen, of these upon as the

It would nonusable, cycles.

desirable

elements the presence

high The (figs.

molecular 5). The

establishing goal the cycles

biogeochemical of building

biological 4 and

cycling lack atmosphere

of elements of oxygen as quickly

of oxygen

an oxygen-containing cycles on which Earth,

possible, lar oxygen.

demands Such

creation do of not

of biological appear

do not

utilize

free molecubecause of the of

to occur

probably in

physiological elements. While efficient, task does in principle. organism, amounts expense not

limitations

the

numerous

organisms

involved

the

cycling

minimizing microbial

the

enormous ecology, of the on

complexity particularly one recycling anaerobic result

of designing not requiring schemes growth in the organic phosphorus

a planet-wide, oxygen, this

steady-state not seem The such of of carbon

to be insuperable. overall effect

Reasonable massive Mars nitrogen, h_, --* C(H20)

do exist,

at least

of a photosynthetic generation compounds and water: of large at the

as a cyanophyte, sulfur, and inorganic dioxide,

would sulfur,

nitrogen,

phosphorus-containing

CO2 where organic oxygen:

+ NO-3 + SO2 + PO4 C(H20) compounds, represents etc. The

+ H20

+ R-NH2

+ R-SH

+ R-PO4

+ 02

carbohydrate, task

R-NH 2 this

represents flow without

nitrogen-containing utilizing molecular

is to reverse

C(H2

O) + R-NH 2 + R-SH

+ R-PO 4 -+ CO 2 + H 20

+ NO_

+ SO_ + PO_

The carbon organic

complete dioxide material

cycling and

of water

carbohydrate aerobic are and liberated. is not

on In

Earth As

depends

upon

a series of these

of oxygen reactions of the

utilizing

reactions

called

respiration.

a consequence environments, dioxide.

anaerobic to carbon

much are,

remains

oxidized

There

however,

47

microorganisms Oz:

that

can oxidize

carbohydrates

to COz utilizing

nitrate

instead

of

C(H20) Most of the organisms

+ NO_ --*CO2 + HzO + (NO_, that can carry out NO_-mediated the recycling the synthesis

N20,

N2). respiration will

anaerobic

utilize oxygen if it is present. Thus anaerobic respiration would necessitate

of carbohydrate on Mars by or discovery of strains unable complex (fig. 4). by of

to use oxygen for respiration but still capable of using NO3. The movement of nitrogen through the biosphere is quite Inorganic the into NH_ nitrogen either process in the form of NH_, derived of nitrogen process fixation, The organic biochemical organic

from atmospheric is released

nitrogen in the form

or NO_ is assimilated nitrogen The released

by organisms

nitrogen

compounds.

by the biochemical

of deamination.

NHg is transformed

into NO3 by various aerobic microbes and re-utilized in the synthesis of cell materials. As mentioned above, NO3 is transformed into N2 by anaerobic respiration. The release of nitrogen be carried from the organic compounds produced by cyanophytes seeded on Mars could (autolysis), out by enzymes released by the dead cyanophytes deaminating organism themselves for this

or by a second,

nonphotosynthetic

purpose. On Earth deamination is accomplished biologically in anaerobic environments by bacteria such as the Clostridia. The microbial transformation of NH_ to NO_ is, on Earth, an aerobic referred to in the discussion nisms NH_ tions. carbon isms organic must for be found: cyanophyte growth, process. The constraints to the use of aerobic processes of carbohydrate recycling suggest that alternate mechacan be toxic. maintain could been organisms tolerable be further The utilization NH_ concentrato organcertain decomposed of however, organic cyanophytes conditions might respiring have

NH_ at high concentrations compounds

The residual dioxide (fermentation). compounds

nitrogen-free Some under

by either

the anaerobic

or the deaminating to assimilate Although

shown

of low light intensity.

such assimilacompounds

tion generally does not lead to growth, it would help in cycling organic and in supporting growth at very low light intensities. The biological form of sulfur that the death of the organisms sulfide teria

cycling of sulfur on Earth is outlined in figure 5. Sulfate, the cells assimilate, is incorporated into organic compounds. Upon cell, the sulfur oxidized is released sulfide, by various putrefactive aerobic conditions decomposing conditions anaerobic sulfur the bacthe hydrogen oxidizing a toxic gas. Under

in the form of hydrogen However,

S= is spontaneously can oxidize

back to SO_. Under anaerobic if light is available that It is possible these

will accumulate.

photosynthetic anaerobic,

S= to SO4.

bacteria might be adapted for growth on Mars. Alternatively, tion of S= can be accomplished by Thiobacillus denitr_Dcans: 48

the anaerobic oxidathis bacterium couples

the

oxidation then by

of enter

S= to the

SO_

with

the

reduction

of NO_ its mass, NH_.

to nitrogen. or could The overall

The flow

nitrogen biological ot" car6.

would fixation bon,

atmosphere,

increasing

undergo

cyanophytes and sulfur

or Clostridi_mz on Mars,

sp. to form above,

nitrogen,

as discussed

is diagrammed

in figure

ATMOSPHERE

i
/ ORGANnC I

"

_' _ I

/_1

_"

MOLECULES /I I
' l

MACRO-/I I _

I OX',GENBODUCERS P I'_

P.OTOTROP.,C I

o'S/
,._5/

_\_1 "_ I _% _4,..,---_%.:_:_._

o.

\%%'%.
N_"v

"'o

/L,
(::3 _I SMALL 1 /

,--L
N' _, BACTERIA

L:J
M I.C,ROMACROI NUTRIENTS

Rcoo.
RCH2NH

MOLECULES I
2 etc.

RCH.O.,

,_--

-1

ORGAN,C / I
e.g, THtOBACILLUS DENITRtFICAN,S SURFACE LAYER

_O7] \1
_

t
H20 | MELTING

t
H20 | PERMAFROST

t
H20 I

Figure 6.

Biological recycling

of carbon,

nitrogen,

and sulfur on Mars.

49

6. SURVIVAL

AND

PHOTOSYNTHESIS MODELS standpoint how and How

OF TERRESTRIAL

ORGANISMS

ON

MARS: COMPUTER From environment. vary during atmospheric would result the biological

FOR LICHENS the study

AND CYANOPHYTES group was interested in examining organism and/or which

several questions

concerning cycle,

the interaction does the how severe

of a terrestrial temperature water

organism with the Martian of a hypothetical to the ground organism

For example, a diurnal temperatures?

closely is the water,

is it coupled low pressures,

loss by the

from the lack of atmospheric

and winds character-

istic of Mars? Can desiccation be significantly limited by varying the resistance properties of the structural components of the organism within reasonable limits? How might mates terrestrial these various environmental production survive could under factors be made? affect photosynthesis, that any Martian and can estipresent day without treatment, that even a of photosynthetic organisms Is it likely

the envisioned require

conditions complex

extensive genetic engineering? Definitive answers to these questions for which there is presently limited

an extremely

information.

It was felt, however,

simplistic model for such processes could yield significant indications of important trends, and could also serve as a starting point for future, more complex treatments. Two similar biological behavior computer such models were formulated placed speed, factors (oxygen and dioxide as solar flux, wind to water and carbon are considered, of photosynthesis of the derivation to predict diurnal certain aspects of Mars. and atmosphere and equilibrium can be predicted. of the model are of the of lichens and cyanophytes on the surface surface movement, evolution) description

When variables temperatures, constants rates of water

resistance

for photosynthesis, loss and rates aspects A.

such as organism temperature,

The quantitative presented in appendix

51

GENERAL Organism

CONSIDERATIONS Temperature the coupling between organisms and the Martian organisms that the environment, an

To determine energy lichens ture balance. balance -

model

representing

two terrestrial assuming

blue-green organism

algae and

has been developed. organism The transpiration physical

Given certain rate can then

environmental be determined

parameters, as well

the temperais in energy as the rate of are:

of the

can be determined

photosynthesis. The major processes controlling the temperature of the organism (1) solar and thermal radiation, (2) convective and conductive energy transport and (3) evaporative cooling. Solar and thermal radiation from the atmosphere provide an energy source for the organism: the solar radiation reaching the organism upon the transparency of the atmosphere and varies with the Martian season and atmospheric gas which absorbs surfaces Heat time of day. temperature, can absorb transport if there The planetary primarily the radiation or thermal radiation lowest few kilometers, For all practical upon it. occurs at the upper the organism and lower and the air between depends latitude, the of

depends upon and the amount

this thermal either

radiation. incident

purposes,

the organism

all of the planetary

to or from

the organism

is a difference

in temperature

and/or ground. At the upper surface molecular conduction transports heat down the temperature gradient and across a laminar boundary layer. If the air temperature near the ducted transport lower ences than surface from is lower than the temperature the boundary of the organism, layer and rapidly by conduction which occurs occurs if the organism's with the surface cooling solid and then heat diffused is conby eddy is the organism through

into the atmosphere. the air temperature. with the surface.

The opposite Heat transport Molecular emission

temperature if the organism strongly

is in contact rid itself

conduction Two processes and

influto

the temperature of energy

of the organism. are thermal

allow the organism (transpiration). liquid

evaporative

Every body radiates energy according radiate nearly as much as black bodies. Photosynthesis

to its temperature;

bodies

Of critical importance is the potential for photosynthesis of any organism on the Martian surface. It is acknowledged that photosynthesis complex elements. complete, represent variables, set of metabolic The consideration is far beyond the organism's which include interactions, the scope which are subject model. to numerous The study of photosynthesis of the present on this level, although

terrestrial involves a controlling more to

perhaps group

chose

photosynthetic rate as a function of several major limiting temperature, light intensity, carbon dioxide concentration, 52

andwatervaporconcentration saturation the organism. temperatureange or of The r over which the hypotheticalorganismmay carry out photosynthesis limited to is that of approximately260-285 K. Photosynthesis, all cellularmetabolicprolike cesses, sensitiveto freezing;temperature is ahd waterlossgenerallyprovidethe controllinginfluences photosynthesis the model. on in Resistance WaterTranspiration to andCarbonDioxide
Diffusion dioxide are rate, and diffusion diffusion deteris air

The diffusive resistance of the organism to water vapor and carbon critical model parameters since they influence water flux, photosynthetic the space temperature of the organism. cell layer The total resistance to water The carbon vapor dioxide equivalent resistance to the sum of the includes resistance of the cell wall covering,

the intercellular

and the desiccated

(for cyanophytes). All of these values

all of the above and, in addition, the cytoplasm.

the water-saturated have been

cell wall, the

cell membrane and mined or estimated. LICHEN The

empirically

SCHEMATIC schematic indicates, sand. representation the lichen The segment upper of the lichen is envisioned It could model is presented in figure 7. As surface, and the surface, of in a

the diagram covered with

as growing which

on the Martian just beneath is modelled

is fully exposed parallelepiped this segment

to solar radiation. of square

also be buried

of the lichen

is a rectangular symbionts

surface,

with an edge length

of 1 cm. The thickness

is T cm, including

the layer of active

algal and fungal

homogeneous matrix (25% cells by volume), and a very thin "cuticose" protective layer over the top surface. Its total thickness can vary from 0.05 to 0.3 cm. A non-turbulent air layer presents Inputs an additional to and outputs boundary assumptions to diffusion, and is a function of the wind speed. in the cyanophyte cyanophyte model, photosynthetically perature 1972). from the system are the same as those

model.

The basic general

are also the same as in the

except that the minimum temperature at which the organism is active is set at 260 K (as opposed to 273 K). This cut-off temin light of published observations (e.g., Lange and Kappen,

is reasonable

CYANOPHYTE Figure

SCHEMATIC the proposed schematic for the blue-green algal model. The

8 shows

organisms are envisioned as existing may be surrounded by a gelatinous surface of the planet.

in a thin layer of interdigitating filaments. They covering and can reside at or just below the crusty or "cuticose" in

Since such algal mats are commonly 53

/ 0

C:}., Lu

B.
@

_.)

_-_

o_

_B
= &

22
l

N
e_

"N
i-_-z G0(J 0.,_

54

E E
X

\ \ \ \\ \\

,,)

+.:,
0 e_ 0 m

_..._

I.

55

their uppersegments, desiccated a layerisassumed overliethe "active"cells.This to desiccated layermayinitially resultfromexposure cellsto therapidmoistureloss of which would be characteristic the Martianenvironment.t is assumedhat this of I t layermay attain a thickness equivalent o that of theactivecelllayer,andthat this t desiccated matrix provides anadditionalbarrierto waterlossby the underlyingcells and probablyprotects from ultravioletradiation.The resistance propertyof the desiccated layer is an importantvariablein the model.The thickness the active of cellshasarbitrarily beenchosen 500_m. By terrestrial tandardshis is relatively at s t thin, however,t is assumed i thatthe severe Martianenvironmental conditions would not permit luxurient growth.This layer is in intimatecontactwith the surfaceor immediatesubsurface f Marsandconsists o arbitrarilyof 50%cellsby vohlme.The remainderis proposedto be water-saturatedir. In view of the extremelylow a atmospheric presstires n Mars,it wasthought unreasonable postulatea liquid o to waterlayer,althoughits presence ould be of little consequence this model.As w to statedearlier,the thickness andresistivity(resistance er unit thickness)was aried p v to establish theinfluenceof this propertyuponcertainof theotherparameters.
RESULTS The environmental models of blue-green algae and lichen discussed in the pre-

vious section have been applied to Martian conditions to determine if these terrestrial organisms could survive the harsh Martian environment, and if so, whether they could bring about any appreciable change in atmospheric composition by photosynthesis. and The two crucial assumptions made are that that the organism can withstand the intense there polar may well be large amounts caps. If so, then the possibility then any biological modification water is available to the organism ultraviolet radiation. As discussed ice locked that in the regolith and this ice could be released Regard-

earlier remnant

of water exists

and utilized, ing ultraviolet the algal matt: discussed.

at least on a local scale, for the growth radiation, the desiccated layer the lichens have a high tolerance

of the organism;

clearly if water shielding for as previously

is not available

of Mars appears

impossible.

could provide adequate to ultraviolet radiation,

The model study has not been compared with laboratory or field situations; in most cases these data are not available. Nevertheless, the results for these models are presented since they provide insight to the importance of various physical processes coupling the organism to the environment and may suggest future laboratory studies which will allow the model to be refined The equatorial particular equinox Martian situation environment which and used for predictions. chosen should 56 be for the most model corresponds to to an conducive terrestrial

organisms noon ism and

in terms ly/min.

of temperature The from values back the for

and

radiation. from for ly/min. wind are to the the the

The

maximum

solar

flux on the

at local organeffect and might with

is 0.86 was

radiation calculations 0.023 near-surface wind

atmosphere carbon speed

incident dioxide

determined different Martian

greenhouse used: low 1,500, values

is considered Three

to be a constant

have

been but

1000 well little

cm/sec. represent irregularity. The

surface regions

velocities

uncertain, highest

sheltered

as opposed

velocities

at surfaces

ground

and

air temperatures while the wind and 2 ). Note thermal important seven though numbered speeds, their that

are

given curves

in figure correspond

9 by to

the the

upper

and

lower algal i
z 2 2

curves

respectively, for (b,

calculated

temperature layer tions surface plant surface ture of the that layer, and the perature because sphere. and the the day. of thickness resistivities

different (a, the most cm) min/cm

desiccated WJNDSPEEP I-_- ] " h

i , I II 500 1000 500 1 crn/sec [ 0 cm ! 0 min/cm 3 rnin,tcm; crn,_sec f 0 03 cm I cm/sec cm/sec i III t IV I V _ O03cm 3 minJcm f 0_05 cm ! 333 min/cm iO.05cmL333min/cm_

corresponding for all condiwith and the the

considered, is the

conduction factor, degrees

lO00cm/sec

LOCAL

TIME, 1200

hr 1400 I 1600 I 1800

temperature temperature, near the ground

is within even

of the

0600 300

0800 I

1000 I

the air temperaTEM E

is some is no The mean with heat that of

65 K less than desiccated of the the layer organism

that (prois

290

surface; the the the

if there surface.

280

file 1) then

temperature

thicker thermal will

desiccated

270

lower lower also the

the

conductivity be. The temspeed atmospeed wind to the of wind layer,

a. 260

temperature

decreases

increased transport

260

of convective Also note

__.240

regardless the 273 can

230

condition will Thus be on blue-green variation the in first second is the day being the

desiccated K about of the be used

the
220

organism temperatures time the The mat layer, and the Each been during the the layer

above Mars

6 hr during surface the to predict

a knowledge

diurnal

210

algae in the

will be above water loss for

freezing. an algal 10. the The layer of

2O0 0600

0800

1000 LOCAL

1200 TIME,

1400 hr

1600

1800

is shown refer the resistivity;

in figure to the the of

Figure

9.-

Diurnal

variation

of

numbers

parentheses

desiccated

thickness

resistance two numbers.

product layer with

of these and

blue-green algae temperature for Martian equatorial equinox conditions. Algae thickness is 0.5 mm, a is the thickness of the desiccated layer, and b is the corresponding resistivity. Profile I overlies the ground temperature profile.

desiccated combined ranges

thickness the three

resistivity speeds;

has the

wind

resulting

correspond

to the

width

of each 57

10 2

i I0,0)

101

100

10.1

10_2

io 3

:=

104

10% WATER FREEZING

LOSS POINT 1200 TIME, I 1100 hr

i0._c_:_ / 0700

0800

0900

1000

1100 LOCAL

1000

0900

0_00

0700

0600

Figure

10.-

Diurnal

variation Symbols

of transpiration are explained

rate in text.

for blue-green

algae.

shaded

region.

The upper

boundary boundary

of each shaded a speed

region refers to a wind speed of The solid arrows on the

1 cm/sec,

and the lower

of 1000 cm/sec.

uppermost three curves are the times for which the organism has reached a 10% water loss, the total amount of water in the organism being 25 mg/cm 2. For the lowermost reach two curves which reaches represent the largest as indicated. 273 K and through does not resistivity, is assumed ceases when is less than occur the water loss does not are the times for water 0.15 before after content (uppermost the freezing subsequent is to occur only when an the about even 10% but is either temperature by curves), 10%. Notice so that 1 or 2-1/2%, has reached that The open arrows

which the organism organism's reduced three point

273 K. Photosynthesis if the resistance

a 10% water

loss is reached photosynthesis

sublimation

is reached,

warming. If no desiccated layer is present water loss would be complete and the algal mat could not survive. A desiccated layer, with resistance greater than about ten, appears being wind necessary the resistance for survival of wind of the organism. speed on the transpiration layer. Contrary rate is small, the major factor to expectation, the larger the region). cooling (each shaded for evaporative reduces The influence

of the desiccated

speed, the lower the transpiration loss, nevertheless

rate for each resistance a smaller resistance temperature

While the larger wind speed does produce and water saturation resistance

the lower organism

the water vapor

pressure within the algal system which overcompensates for the smaller and gives a lower water loss (appendix A, eq. (AI4)). Note that the wind in water loss; the diurnal variation in water loss, primarily

speed is not a major factor

58

controlledby surfacetemperature, several rdersof magnitude, hilea change is o w in wind speed changes waterlossby lessthana factorof two. the The rate of photosynthesis essentially is constantoverthe day for all models considered. Thereasonfor this is that the ratecoefficientfor radiationis verysmall, being1.210 ly/min, so that evenat 0700hourslocaltime,whenthe solarradia-3 tion is 0.2,the ratio 1.210 -3/0.2 << 1,andthe photosynthesis rateis independent of light intensity. The numbersby the open arrowsof the lower two curvesof figure10 refer to the molesof carbondioxidefixed/cm 2/day,andorganisms ith w desiccated layersof resistances greaterhanabouttenwouldyield about10 moles t -6 of oxygen/cm 2/day. If, for example, one-fourth of the planetary surface were covered by blue-green algae, and if they were able to photosynthesize for about half a Martian year, then in seven thousand Earth years, an amount of oxygen would have been produced equivalent to the present amount of carbon dioxide in the Martian atmosphere, Similar that is, 5 mb. calculations have been carried out for the lichen model. The same

atmospheric model has been assumed, that is, equatorial equinox conditions and three different wind speeds; 1, 500, and 1000 cm/sec. Three different lichen thicknesses were used: Diurnal speeds are shown 3 mm, 1.5 mm, and 0.5 mm. in the lichen 11. For temperatures sheltered for different conditions, thicknesses that is, very and wind low wind in figure variations

speeds, the lichen temperature is closely coupled to the surface temperature, even for a thickness of 3 mm. This occurs because the thermal conduction to the atmosphere becomes the for is small. larger; As the wind speed increases, the heat loss to the atmosphere from also the thicker the organism, the smaller is the heat transport

surface. Both effects tend to decrease the organism temperature. a lichen thickness of 3 mm, and wind speed of 1000 cm/sec, of the lichen is only 262 K, although and photosynthesis. that is, with a wind of 1 cm/sec, the ground from figure conditions, 11 that wind speed is closely coupled

For example, the maximum is 300 K. to the organism loss is large

temperature and influences

temperature

Thus it is apparent For sheltered

both transpiration

the water

and is not strongly dependent on the lichen thickness (fig. 12). Note that for a given wind speed, the greater the thickness of the lichen, the smaller is the water loss. This is a result of the lower organism thickness. It is also apparent from water as that organism loss decreases temperature with increasing which lowers for the algal model temperature associated with the larger lichen figure 12 that for a given lichen thickness, the wind speed. the saturation The explanation water vapor for this is the same a lower layer pressure and thus the is, the larger wind speed produces in air boundary

(fig. 10), that

water loss; this process is more important than the decrease thickenss which decreases the resistance to water loss.

59

LOCAL 0600 300 F 0800 1000

TIME 1200

hr 1400 1600 I 1800 | 0800 1 1 cm,_sec 500 1000 =_ E cm,lsec cm,'sec 1000 '. -----03 0.05 1200 1400 i 1600 I

i
290 t I _

A
B

1o2 [
i
C

280

270

E 101 2600.05

g
i-< 250

g
_ 240

230 TEMPERATURI 220

I00_

210

0 05 O3 (on1) 005

!
J
I ! , 1800 10 1 L 0600 J 0800 1000 LOCAL 1200 TIME, 1 1400 hr 1600 1800

200 _
E 1!]0 [ 0600 L 1000LOCAL 0800

_oo
I000 1000 i 1400 hr 1200 TIME,

o3
005 03 1 1600

Figure lichen

11.-

Diurnal

variation for

of

temperature equinox

Martian

Figure tion

12. rate

Diurnal variation for lichen. Numbers of the lichen

of transpirarefer to the

equatorial

conditions.

thicknesses

layers.

Figure speeds

13 shows

the rate of photosynthesis The photosynthetic

(moles CO2 fixed) for various rate is less than half that

wind of the

and lichen

thicknesses.

blue green algae due to the larger rate coefficient (K2). Note that the diurnal variation of photosynthesis exhibits the same relationship to wind speed and lichen thickness as does the transpiration rate: that is, as the wind speed or lichen thickness increases, the transpiration sis occurs. This can most synthetic the product rate (appendix Ar' decreases, At' decreases while a corresponding increase in photosyntheeasily be explained by referring to the equation for photoA, eq. (A12)) where, + K3)/(S20)[ri(Dtt20/Dc02 ) + 61 and noting that photosynthesis increases as

=- (1 + K2/1)(1

and 6 cc V _,,2. The terms 6 decreases. water photosynthetic Also a larger S. Both rate. retention

are explained wind speed parameters

in appendix means

A. As the wind speed increases, water loss and thus a larger At' and increase the

a lower

therefore

tend to reduce

6O

14 r

l
_ _ 10_ 053 _

oc

0600

0800

1000 LOCAL

1200 TIME,

1400 hr

1600

1800

Figure 13.Diurnal variation of photosynthetic rate for lichen. Numbers refer to the thicknesses of the lichen layers. An additional factor is the of 260 released thicknesses. moles/cm algae. about equivalent slow growing produce because could At 70,000 to and time which the must be taken into account to determine point the daily lichen; fixed wind is of for by of are that

photosynthesis a temperature or oxygen and to 10 -v

temperature and the

is above number per day

the freezing of moles

of the dioxide various the

K is assumed per square The

of carbon for that less

centimeter are hours shown per one

is calculated 19. Note an

speeds able about

results

in table day order of the be of yielding

organism that

photosynthesize

several

oxygen

production than

2 , approximately this rate, Earth the any if one-fourth years

of magnitude Martian surface

blue-green lichens, oxygen very lichens However, in generating

were an

covered amount lichens appear

would amount change

required carbon

to produce dioxide. Thus

present significant tolerance

Clearly, not lnight concentration they

such

a coverage

is unrealistic. in the

it does

oxygen radiation,

on Mars. be effective

of their

to ultraviolet

an ozone

shield.

61

TABLE19.- OXYGEN PRODUCTION LICHEN BY

THICKNESSES Local time when water loss reaches

FOR AND WIND SPEEDS

VARIOUS

LICHEN

Thickness

(cm)
0.3 0.3 0.3 0.15 0.15 0.15 0.05 0.05 0.05 Note: Martian conditions.

Wind speed (cm/sec)

1.

Time above 260 K (local time) 0925-1623 1124-1500 1200-1413 0927-1625 1030-1600 1100-1530 0930-1628 0947-1618 0953-1613 and radiation

Hours available for photosynthesis 3h35. m 3 h 36 m 2h 13 m

80%
1300

Oxygen released (moles/cm 2) 3Xlff 7

500. 1000. 1. 500. 1000. 1. 500. t000. temperatures

4Xlff 7 3Xlff 3Xlff 5Xlff 5Xlff 3Xlff 3Xlff 4Xlff equinox 7 7 7 7 7 7 7

1300 1700
...

3h33 m 5 h 30 m 4 h 30 m 3 h 30 m 4 h 13 m 4h 7m

1300 1400 1400 field correspond

to equatorial

62

7.

PLANETARY ENVIRONMENT From previous

ENGINEERING:

MODIFICATION

OF

THE

MARTIAN

considerations

it is clear

that

modification

of the

Martian

environment is necessary if extensive growth of terrestrial organisms is to occur in a reasonable time. A consideration of the evolution of the terrestrial planets indicates that the modification of the Martian environment to similarity with that of Earth is possible. Although the evolution of planetary atmospheres is still a controversial topic one, years (Cloud, 1965), it is generally agreed that the Earth's atmosphere is a secondary having outgassed from the lithosphere over a period of approximately 4 billion

(Berkner and Marshall, 1965). In view of the similarity in the material from which the terrestrial planets were formed (Pollack, 1975), it is reasonable to assume that their atmospheres are also secondary is possible to construct a hypothetical for Venus, ently Earth, and Mars (Rasool, in nature. With this essential point assumed, it but plausible history of atmospheric evolution 1971), and to demonstrate that Mars is presof the three atmo-

in a state appropriate for atmospheric modification. Calculation of the initial, mean planetary surface planets, yields Venus, Earth, and Mars, prior temperatures of 350 K, 270 K, and

temperature

terrestrial sphere,

to an outgassed

secondary

220 K respectively.

As volcanic

gas, primarily carbon dioxide and water, accumulated on Venus and Earth, which were initially warm, a substantial greenhouse effect _ developed. In the case of Venus the _-500 temperature increment vapor of never the greenhouse effect was enormous (presently K), since water On Earth greenhouse condensed into liquid form due to the high initial forming the oceans and limiting the K). Moreover, water in liquid form

temperature. incremental

water vapor condensed, effect (presently 237

Atmospheric greenhouse effect refers to a condition of increased warming caused by the presence of certain gases in the atmosphere. These gases, carbon dioxide and water vapor among them, absorb some of the thermal radiation (heat) rising from the surface of a planet, so that not all of the radiation is lost to space but is re-emitted down from the atmosphere thereby heating the planet's surface. 63

aided

in the

reaction Significant

of

carbon

dioxide of carbon the

with dioxide

silicates also

in the dissolved atmosphere down the

lithosphere ill the oceans

to form in the due

carbonates. form to the evolution On ture and water 24 from K). outgassing.

amounts On Venus,

of bicarbonates. absence of liquid

carbon

dioxide Venus

is still present path of the

water.

In order to cool was entirely and carbon of present

to guide the planet different. dioxide carbon amount

Earth's

it would Mars the the

be necessary situation vapor large hence for the lack

significantly. The low and initial into water surface ice after are present carbon effect dioxide temperavolcanic as ice and

caused

water and

to crystalize dioxide of atmospheric greenhouse the

Presently,

amounts the

permafrost, is

insufficient dioxide Mars moderate form,

a measurable of liquid

incremental water prevented

(presently

Moreover,

formation

of carbonates

carbon Evidently,

and silicates. has yet to follow the path on of Mars Earth's would evolution. be required Unlike on

Venus, water

only in liquid

temperature the necessary

increases catalyst

to provide

for Earth-like

planetary

evolution.

INCREASING

MARTIAN

SURFACE

TEMPERATURE

It is obvious extensive may can require An be gained (Ward rely biological insight by

that

an increase are

in the to

surface and, which

temperature indeed, might even affect long Sagan the bar. solar

of Mars their

is necessary mere existence change climate These

if

processes various the

occur,

a temperature to the reviewing 1974;

modification. mechanisms several a temperature term etal., resulting sublimation absorbed Martian 1973). papers and of the up concerning 1973; cap one

changes mechanisms increase cap poles. energy could

etal., on

Gierasch ranging by cap,

Toon, polar to about

a sublimation mass about the polar

with

substantial of the by the of solar or orbital of the of the until the

in atmospheric be brought increased on of Mars, additional result warming This in the

The

an increase could for example, solar

in available by

energy the

This

absorbance

be caused output, to (see

increasing

the amount obliquity the albedo

incident

by modifying

parameters caps. planet effects. supply As would

by an increased carbon both would dioxide from accelerate

or by decreasing the the atmosphere, next of carbon

is added advective the

a warming dioxide

section)and

greenhouse

sublimation

cap is exhausted. significance carbon They on concluded this of a remnant mass that was the Ward carbon initially polar dioxide ice cap to the control by controlled Leighton the of and atmo-

The the Murray spheric in total tions.

possible (1966).

atmospheric

dioxide

investigated determined for various to

CO 2 ice cap

CO 2. Based atmospheric They find

assumption and polar

et al. (1974)

the variations obliquity maximum oscillaand

pressure that the

cap temperature extremes 64

climatic

corresponding

minimum 130 water K to ice. to about

obliquity 160 K with 30 mb.

would Their

cause

the also

remnant pressure indicates

polar that

cap the

temperatures of a few south residual

to range tenths cap should

from be

a corresponding analysis

variation

of ambient

Ingersoll carbon partial Also, dioxide pressure. the

(1974) reservoir Recent

has

argued in the observations heat

against polar caps

the

assumption which residual the due

that

there polar

is a permanent the cap atmospheric is water might lead ice. to

of Mars

controls

indicate transport

increased that

poleward

to increasing cap. to change advective, cap ice, carbon cap remnants then

pressure

an instability, It appears ture polar and caps

is, a continued that initiate the is by might is both remnant

sublimation the only plausible

of the way

therefore

the surface sublimate greenhouse, and the

temperaof the or both. amount can of

pressure which concern

a mechanically a "runaway composition were carbon

induced effect"; of the dioxide buried

reduction

in the

Of major

ice available. more no of degrees, carbon the than

If the a few for dioxide surface

its thickness would then _8 kg,

be no and eight to possiof kg, has be with be at about of need from is the is

kilometers, this has

otherwise been thickness to found. of the

the three

dioxide semidiameter is assumed, 3X10 There and

liquify,

evidence with

If a polar atmosphere one

of about enough

an average

kilometers,

the amount

available atmospheric more dioxide a surface between in the

is about hundred fold.

increase bility the

pressure carbon the

is also the estimates

1019

of a buried, total The carbon yield relationship discussed

extensive in both pressure literature

dioxide

reservoir, and crust

recent

atmosphere of several ice cap regolith. and partial to be bars and recent

suggest

about

which not

would

if all could a runaway evidence to have

be released. greenhouse suggests effect there may

a water ice in the

been

although

as much liquid-water, least 273

as 10 _ 9 kg of water the K and of vapor. 300 increase dioxide effect (Noll the rate, surface 6.1 mb that

In order pressure present,

an equilibrium vapor water, in the be must or

temperature respectively. thought

of water 1016 kg of must

Approximately

one-thousandth atmospheric to be only The a carbon greenhouse ture added adiabatic lower the the profile to curve. ground ground.

possibly cap with surface

form

A remnant to provide global in mean greenhouse if the and McEIroy, The

semi-diameter of vapor. temperature in figure rate

of eight which might

degrees

would

m thick

this quantity

be expected line

effect

is shown lapse

14. The uppermost to the mean as additional temperature, effect and carbon given corresponds

temperature 1974) most

corresponding

temperadioxide by

is maintained rapidly influence twelve varying

atmosphere. represents The dashed

a dry to the

a maximum line shows averaging the about

greenhouse

of the discontinuity degrees above B. the

at the

surface: at

temperature Details

air temperature

of the calculations

are given

in appendix

65

I000 I ! I 1 | I I / I I

If all the estimated

carbon the of

dioxide result-

MEAN

GR Hg SE /
> ua .a 100

on ing

Mars

could

be volatilized, pressure bars would

atmospheric three

approxithe

mately

increase

-'I
n .01

mean surface 10 and 20K. amount dioxide of ice, gas,

temperature To release if it were require all

between such an carbon kcal;

would

102_

ua

an amount of energy equivalent to the total radiation incident on Mars for three Martian years. that that and a runaway effect

//..
0

/_

GREENHOUSE

It is possible could cally

| 1 I 2 I 3 l 4 I 5 I 6 _i 7 J 8 K i 9 I 10 11 .001

be small

induced; increase increase

is, a mechanidistribdioxide

generated

uniformly in carbon the

uted which and allow Such

TEMPERATURE

INCREASE.

would

temperature

Figure 14.- Carbon dioxide greenhouse effect. Uppermost line refers to a mean temperature profile while the lower line is for an adiabatic lapse rate. Heavy line refers to the slope of the sublimation equilibrium surface for carbon dioxide.

partial pressure sufficiently to continued sublimation to occur. a process requires that

AT Apgh.

>

dT dPequil.

that less the the

is, the than

slope the ratio

of the equilibrium of the temperature increase for various from the in

curve carbon

for carbon dioxide effect

dioxide

ice and

vapor

should effect of

be to the

increase

produced partial is given

by the greenhouse pressure. is shown in figure The 14. For slope

corresponding curve slope AT/Ap

equilibrium

temperatures greenhouse

and pressures

in table

20,

while

an instability

TABLE 20.-

TEMPERATURE, PRESSURE, AND SLOPE FOR EQUILIBRIUM BETWEEN CARBON DIOXIDE ICE AND VAPOR

Temperature, K 140 160 165 170 180 200 217

Pressure, mbar 1.14 18.1 32.3 56.4 156. 873. 5180.

dT/dPequil, K/mbar 5.6 .46 .27 .17 .067 .0147 .0029

66

to develop, that, initially, over the planet are greater pole would dioxide inequality ide added the than effect. except As supplied dioxide increases carbon

AT/Apgh.>dT/dPequil" sublimation would where about sink. surface 165 K. The Even if the

so proceed winter above

cc

I
l

_ i

w I _l
_

WATER

temperatures

than vapor

be excluded

and act as a carbon

is not obeyed, the carbon dioxto the atmosphere will remain if dioxide induced 15 vapor by pressure the this is less for the true vapor pressure shows

the equilibrium Figure

100 SOLID CO 2 \

CO

GAS

__ WATER

temperature

greenhouse to be

T"'PLELO;NL 2,'K
1000 80 1 100 I 120 1 140 I 160 I 180 _ I 200 I 220 I 240 'K I 260 I 280 I 300 320

over the winter pole. carbon dioxide is continuously to more the atmosphere partial the carbon pressure Figure 15.- Pressure-temperature cross section of the thermodynamic surface for carbon dioxide and water. temperatures. is doubtful; even and could pole For example, the carbon be released if the entire dioxide vapor vapor
TEMPERATURE,

equilibrium slowly

than the correspond-

ing pressure effect, and only occur polar pressure pressure sphere would

increase from the greenhouse continued sublimation would from increasingly were carbon 5 mb over a bar could higher the surface summer dioxide (which

cap remnant is about of about

though

an equilibrium

be maintained)

to the atmo-

- an increase of about two hundred fold - then the mean global temperature increase about 7 K and the pressure would be about 500 mb. Continued could would only occur not take dioxide. similar when temperatures place since were above about could 210 K, however, more than caps the atmosphere support

sublimation redeposition

one bar of carbon are not carbon vapor An analysis

Such analysis to that

also suggests strongly dioxide has been

that the residual carried than

dioxide

but water temperature

ice (fig. 15). for carbon increase out for water that for an is significantly greater

(fig. 16). The

equivalent amount of carbon dioxide. For example, if enough added to the atmosphere to increase the total pressure by effect carbon The dioxide in the one would dioxide reasons increase would that the temperature change about 10 K, whereas effect the temperature vapor greenhouse by only about is greater

water vapor could be 10%, the greenhouse amount of one-tenth than that of a degree. for carbon

an equivalent

the water

are twofold: while the carbon dioxide absorbs planetary radiation primarily 15 #m spectral region, water vapor absorbs strongly in two spectral regions, near down 6.3/am to about and the other Water 67 centered vapor about 80/am with significant in the 18/am. also absorbs solar radiation

centered

absorption

101

] .i / ./

/ L: E _d / 10 0 / / / / / / / /

,_

c:" 10 1

_2

-->

-- --

gnls/cm VAPOR

2 COLUMN PRESSURE, mh

10 2

101

10 3 0 2

J 4

10

12

14

16

18

20 K

22

24

26

TEMPERATURE

INCREASE,

Figure 16.- Water vapor grem_house effect. The dashed and solid curves refer to the water vapor amount in units of partial pressure (rob) and colunm density (g/cnl 2 ) respectively. I leaw curve is the slope of the sublimation equilibrium surface for water vapor.

near solar sion tion

infrared, radiation to space are also The

between will and given

about

0.8

_u]]] and and must

6/am.

With

increasing by

water

vapor,

more emiscalcula-

be absorbed thus a higher

be balanced temperature.

a greater The details

planetary of this

atmospheric B. vapor some

in appendix of the

slopes and

ice-water are shown

equilibrium 21. insight could These into

surface values, the

for when

several of

values

of with

pressure the greenhouse

temperature profile effect.

in table vapor

compared into

AT/Ap

in figure For example,

16 give

instability be distributed

a forced the

if water

initially would about

atmosphere I AT/Alhql regions where the surface that would equals is, sublimation occur that l'or that AT/Apgh. would

500 K/mbh temperature continue

then an instability is greater than until a new where

result only for those 220 K (see table 21). is established, which curve the

equilibrium the slope of

surface induced

temperature

the sublimation is less than

by' the greenhouse 68

effect.

If AT/AI}gh.

TABLE 21.- TEMPERATURE, PRESSURE, AND

ICE-WATER VAPOR

CORRESPONDING SUBLIMATION SURFACE Pressure, mbar dT/dPEquil, K/mbar 1.54X 1011 6.13107 2.42 10 s 3.92 103 164. 13. 2.

SLOPE

OF

Temperature, K 125 150 175 200 225 250 273

1.66X 5.96X 2.07X 1.66 5.03X 7.7 IX 6.11

10-11 1 fib 10-s 10-3 10-2 10-1

ice-vapor with the

equilibrium surface from they above could of the would would sublimation climate further large

slope,

and

if the any

water water

vapor vapor are of

pressure

is initially

in equilibrium would primarily imagine, surface cap and Some to the increase, albedo, a to be

temperature, a "mean give

then global

mechanically limited value,

released and one with are might the polar ice ejection in the are

redeposited. Results useful based in that on the

model"

insight that

to physical water

processes. vapor

However,

analysis, be likely be

in equilibrium from pole a remnant

temperature because deposition atmosphere additional permanent investigate

mechanically systems, occur necessary. through over

ejected distributed the winter

water the

wind

throughout and

atmosphere.

continued

With

a greenhouse-induced as well

temperature

instabilities, occur.

as a decrease models formation.

change this effect

might

Two-dimensional cloud

necessary

as well

as probable

ADVECTIVE "Advection" of The tropics turbulent, disturbances tion and air.

TRANSPORT is the on term the used other by meteorologists hand, is usually efficient Warm poleward down to denote reserved horizontal for vertical heat rising transport transport. from the

"Convection," Earth's poles bubbles (advection), turbulent between weather to air

present to the

atmosphere by these and two

is particularly processes. brought is carried

in transferring air, by to the polar

equatorial

in buoyant, synoptic by radiaits peak

(convection), is finally

mid-latitude surface reaches

mixing

(convection).

Poleward

heat

advection

strength baroclinic from

latitudes 40 and 50 in each disturbances are most intense. pole tends mean to reduce meridional Because 69

hemisphere during Such a redistribution temperature of a thin

winter when of energy without

equator the

gradients

changing

planet's

temperature.

atmosphere,

atmospheric

heattransferon Marsis minimalat present, ut it wouldbeof increasingignificance b s if the atmosphericensityincreased. d Usingan energybalanceapproacha crudeestimatecanbederivedof annual heatadvectionat the Martianpoles.Figure17 illustratesthe averagennualenergy a fluxes at the NPC(north polar cap).To facilitatewriting the meanannualenergy balance equations, is assumed it that: (a) the atmosphere transparent solarradiation, is to (b) surface emissivity andabsorptivity infraredradiationareboth unity, for (c) G = 0 over an annual cycle,
(d) ture, polar mean annual polar atmosphere may be characterized by a single temperaTa, and, (e) the surface With may system is in a steady-state temperature) remain fixed. simplifications the equilibrium energy so that balance Ta and equations TO (mean over annual the pole

these

annual

be written:

V + aL o - 2L a = +H+E

(atmosphere)

F + L a - L o = -T-H_-E (surface)

V_

o
l = Q E

I
1

ATMOSPHERE

I'

i
F L a Lo i G

POLAR

SURFACE

SUBSURFACE

Figure 17.Average annual energy north polar cap; V=poleward F=solar radiation flux absorbed

flux diagram for heat advection; at the surface;

L a =infrared radiation lost by the atmosphere; L o = infrared radiation lost by the surface; a = atmospheric absorptivity for infrared radiation; H = sensible heat convection; E=latent heat convection; G = subsurface heat conduction.

70

(see fig. 17 caption the poleward

for definition (V), V=

of terms).

Adding

and rearranging

terms

yields for

heat advection

L a + (1 -a)L

o-F make the

(13) parameterizations: is the Stefandiscussed in a

To

evaluate

the

right-hand

side of equation

(14)we

L o = oTo 4 and L a = KL o, which together imply a = K(To/Ta)4;o Boltzmann constant; and K is an empirically determined constant previous section (p. 21). The factor (To/Ta)4 is a measure effect at the polar cap and has a value near unity for current greenhouse this ratio becomes V = oTo4 -F Equation between the surface (14) yields rate mean annual heat advection quantities. effect). To study advective effects independent is maintained at unity for all calculations.

of the greenhouse conditions (i.e., no effect, (13) equation

of the greenhouse

By substitution,

(14) at the pole as a small difference This can produce a 5 K error poleward heat a large error annual advection. in in mean polar As a

two large imperfectly of advection. will cause temperature

known

calculated

For example, a 25% error with equation

in the

result, estimates

of heat advection

(14) must be regarded cautiously.

Substituting T o = 160 K and F= 40 ly/day (lly = l cal/cm2), values based on table 1, V = 39 ly/day. Leovy and Mintz (1969) estimate atmospheric energy transport during winter solstice across 43.5 N as 19X202 erg/sec. If distributed uniformly over the planet's surface poleward of 43.5 N, this is equivalent to 19 ly/day. The assumption of uniform energy distribution, however, produces too low by at least a factor of 2 for the Earth's north pole radiational same cooling is offset by strong Leovy advective and flux from situation occurs on Mars, Mintz' the estimate values which are where extensive If the

lower latitudes.

will also be too low, advection on the

perhaps also by a factor of two. Gierasch and Toon (1973) Martian radiation sphere ward. carbon dioxide in turn, equator north polar at the cap, and The winter

explored

effects

of heat

cap. They suggested that a 20% increase in absorbed solar if maintained for about 100 yr, might lead to a 1 bar atmowarmer solar capacity climate. flux at the The physical very pole north mechanism close will sublime in advecting to further is straightforpoint of more carbon has a surface temperature to the frost

a significantly pole Increasing heat

dioxide. from to

the cap, resulting the pole. Increased

in a slightly polar

denser atmosphere. leads

The denser air will, heat from the carbon dioxide

have a larger

and be more efficient heating

71

sublimation, has been atmosphere bon dioxide

and sublimated could

so on. result.

This There in the

feedback If the

process carbon

continues dioxide doubt supply

until whether

the the this

entire enough, quantity large

polar

cap

to vapor.

is large

a 1 bar of cara change. changes.

is considerable polar cap may obtain

contained the Stone's waves,

is sufficient still yield of

to produce significant heat

Nevertheless, Using baroclinic

advective (1972) Gierasch

process and Toon

temperature flux by

parameterization

advective

midlatitude

V = oda(T e - Ta)_ for sure, polar and winter heat advection, where and ce = 1.5 polar 10 -4 cm/sec/K z ; P = surface

_15) pres-

Te, Ta are the equatorial this because assumption, it is primarily

atmospheric by radiative

temperatures, temperature and convective

respectively. as being processes

Following constant

equatorial

atmospheric

is taken

determined

which should for equilibrium on figure

be little modified by climatic changes. It is also assumed that T a = T o conditions in the absence of a greenhouse effect. Table 22 is based paper polar by Gierasch surface value Leovy and of Toon 1.8X104 (1973) for a mean annual solar flux and

1 in the at the

absorbed T e = 200 than that this eddy section. heat

erg/cmZ/sec

(=381y/day),

K. The advection given by either This flux may,

found for the 5 mb pressure of table 22 is smaller and Mintz (1972) or the value discussed earlier in be due about to the 65% fact of the that total equation heat flux (15) includes pole. only Also,

in part, comprises

which

at the

TABLE 22.-

RELATIONSHIP

BETWEEN AIR PRESSURE, AND ADVECTIVE FLUX Advective flux V, ly/day 4.6 7.9 16

POLAR

TEMPERATURE,

Air pressure, P, mbar 5 10 25 50 100 250 500 1000

Temperature To, K 146 150 155 162 171 183 188 190

Low temperature Stable climate

27 23

Transition

32

High temperature Stable climate

Source of data:

Gierasch

and Toon (1973),

figure 1.

72

the assumptionof uniform flux distribution polewardof 45

Thus, these To ture figures from could the easily present be low low by a factor temperature 22, by of three. stable climate move

N has

been

made.

to the

high

temperaincrease outside the that solar the high pole can is be

stable

climate This

as indicated might by the current can 20% means (runaway of 190 be brought gases

in table about

requires importing and

all order

of magnitude from

in pressure. the planet, energy transition temperature increased achieved advective polar pressure available by by

an atmosphere (1973) climate solar

volatilizing from the

bound caps. stable

in the low if the polar

regolith,

or somehow Toon

increasing indicate to the flux at

absorbed

polar

Gierasch

temperature annual

stable the

climate roughly some

occur to and

absorbed erg/cm2/sec for

2.2X104 maintained

(= 46.5 sufficient until there

ly/day).

If this

time,

a self-perpetuating with dioxide the high dioxide. increase in one a

process for stable

advection) K is established, If less than

will continue assuming 1 bar by the total

a 1 bar atmosphere is enough of frozen on how carbon carbon a 20% dioxide is present,

temperature

sublimation. climate and may

of carbon quantity

is determined Gierasch be brought parameters

Sagan, absorbed or more

Toon, flux

(1973) about. listed

have This

speculated

solar of the

can be accomplished

by altering

planetary

below: Value runaway needed for

Planetary parameter Distance Orbital Orbital Polar from Sun

Present value 1.5 AU 0.09 23.9 0.77

advectJon 1.4 AU

eccentricity obliquity cap albedo

0.49 31.0 0.73

Only cation. much small required so,

the of

polar Since these

cap the

albedo average are

seems albedo believed

even of to

remotely nonpolar be

within regions

the of

realm fine

of possible

modifi0.25, and

is approximately particles, the

regions of sand If such instability

composed polar cap

a relatively albedo to the years climate or

admixture 0.73.

or dust an albedo might

over could

the

could for

reduce about

be maintained and a stable

a hundred temperature

advective

be triggered

higher

obtained.

73

8.

GENETIC

ENGINEERING

Preceding oxygen sphere the

sections on

of

this by

report

have

discussed The

the

possibility of

of evolving such ways

an

atmosphere would,

Mars take

photosynthesis. long. alteration would

generation to investigate

an atmoby which One climate the such to

however, for

quite

It is desirable might

time

required previously conducive of presently

atmosphere discussed, to plant available present the

be drastically the present might such Such optimal that

reduced. Martian they factors growth niche

approach, one better to more teristics

be to change Another approach organisms Martian required for

growth.

be to alter

characbe far

photosynthetic or altered

would

adapted

to the light,

climate. for

as resistance and photoetc.,

ultraviolet the

temperature rate of oxygen Earth

range

synthesis, could have "ideal"

absolute

production,

ecological which

selection,

be selected evolved, This Martian but

to fit not

environments

contemporary

organisms

Mars environments. discusses oxygenic strategies, using goals, and prospects of genetic for constructing an organism techniques engineering.

chapter

STRATEGY

Organisms for optimal adapt

and growth to

environments and reproduction Earth the To

evolve

together.

Most

organisms

are niche

specialized and do and not the ones. in all in the of using such developadapted

in a particular do not niches resemble

ecological Mars niches

readily organisms Thus, least if likelihood available organisms ment

others. to of

niches

exactly to Martian

adapted growth be meager. time, Martian by

various generate be in the

Earth terrestrial

will be ill-adapted occurs by organisms possible. available to the design at atmosphere to use manner presently

transplanted it would

organisms

all,

it will

an oxygen very most desirable efficient

photosynthesis capable Constructing or under of Mars

possible

niches

techniques possible. strategy

of genetic There might

engineering

is quite one

are numerous

approaches

organisms;

be the following:

75

1. Describein detail the environmental haracteristics f availableMartian c o niches directobservation by andmeasurement. 2. Describein detail those biologicalcharacteristicsequiredof an "ideal" r Martianorganism adapted theseniches. to 3. Isolate"best" terrestrialorganisms ostcloselyresemblingideal" Martian m " organisms searching Earth nichesmost closely resembling prospective by in the Martianniches. 4. Determinewhich characteristicsf these"best" organismseedmodificao n tion ascompared the projected"ideal" organism. to
5. forming The tate Modify "best" the characteristics organisms into parameters so identified "ideal" organisms. of the of organisms factors current best facing or modified adapted organisms they include: strong dust Martian climate dicto these seeded mean ultraviolet storms. The organisms against light, subsurface hemisphere. planetary effect of liquid pressure diurnal on water in the variations further surface part and many climates. on the surface temperatures irradiation, number of by genetic engineering thereby trans-

environmental

the biological The

characteristics

predominant have been large

environmental discussed diurnal conditions, previously.

of

Mars

Briefly, variations,

below highly niches ably ature above be

freezing, desiccating available quite hot limited water spots,

temperature and by frequent

abrasive

for

occupation and determined

contemporary by might factors include, such

photosynthetic as: protection of visible southern mean

is probultraviolet temperIt may maximum harsh soil dust soil layers

radiation,

availability, etc. Such of permafrost

wind niches

protection,

intensity of the in the

local

for example,

regions

in the mid-latitudes moderate would would 273 have increase an

expected

that (e.g.,

a relatively 5 10 K) There

temperature environmental above more storms. ice total More

ameliorating envelope vapor

factors. layers hours and

be a larger a higher increases water K, and

in the atmosphere, and

permafrost, above temperature such that, optimal to increase oxygen genetic

per day

moderated would pools on terrestrial the total

extreme factors the range considered would

be expected of liquid a large space

to optimize might result be of for

environmental present approach these growth, the and changes that

for example, temperature for many greatly production modification. such periods

water The net

of maximum be and

of the planet time available and

would

organisms.

reproduction,

by transplanted

organisms

reduce

requirement In any

for extreme event, adaptation cycles,

to factors extended

as high

levels

of

ultraviolet and others,

radiation, would be

rectirrent necessary. While group Earth

freeze-thaw

of dormancy

no

terrestrial is "best"

organism fit to grow,

is ideally reproduce, Antarctic 76

adapted and dry

to Martian produce valleys

conditions, on

is there Mars? resemble

which

oxygen most closely

Of all the

environments

probably

the

current Martiansurface.Thesevalleyshavefour major oxygenicphotosynthetic groups: green algae,blue-greenalgae,mosses, nd lichens.Comparing a selected propertiesof thesegroupswith a hypothetical"ideal" Martianorganism, is found it that, while noneareideal,the lichens andtheblue-greenlgae a are"best" (table23).
TABLE 23.BIOLOGICAL CHARACTERISTICS OF SOME TERRESTRIAL AND AN IDEAL MARTIAN ORGANISM Extreme resistance to ultraviolet radiation
No

ORGANISMS

Organism

Requires oxygen

Extreme resistance to drying

Growth rate

Growth habitat

Green algae

Yes

No

Fast (hr)

Soil (surface and subsurface, snow (surface), water Surfaces (rock, tree)

Lichen Moss Blue-green algae Ideal Martian organism

Yes

Yes

Yes

Very slow (yr) Slow (wk) Fast (hr)

Yes No

No No

No

Moist surfaces Soil (surface and subsurface), water Soil (surface and subsurface), water

Yes

No

Yes

Yes

Very fast (min)

All groups, oxygen. the rate

other The latter of

than

that

of the

blue-green

algae,

have

a mandatory however, provide other survival can

requirement they for the can grow maximum algae the times on

for in

organisms

will use oxygen and not Pearson

if available; 1970). produced To

absence

oxygen they

(Stewart will to

of accumulation so that

of photosynthetically utilize ultraviolet short.

oxygen, such that algae full effect is not the from is the The

blue-green than

must lichens the

be modified demonstrate surface soil of

oxygen. radiation The

All groups

a sensitivity Mars and The would thereby very slow

be quite could growth On

blue-green from the

occupy

subsurface ultraviolet the goal of

niches

be shielded rate of the and

of incident with

radiation. rapid

lichens

consistent particular the group general is quoted

oxygen

evolution. algae it is concluded

balance, that "ideal" the

considering especially algae organisms. and the

suitability most fitted

of the to be of this

blue-green engineering, modified group of the

as experimental provide by

material, Martian (1963)

viewpoint

of genetic

blue-green

so as to is summarized author:

ecology

Cameron

following

by permission

77

"In general, and cold and water regions Drouet, climatic

the blue-green

algae occur species

in all parts of the world may be distributed

where light from (Gerdel

are available. such 1960), as the

Individual are found Antarctic

in the various

zones, but others

at extreme

limits of the environment, of Greenland of the Dead

or in the cryoconite

and from

the low elevation

Sea to mountains

over 14,000 feet in altitude. 1960), occur in extremely waters Feh6r, (Palmer, 1939). 1959) Planktonic and

They are a part of the salt marsh flora (Chapman, saline Great Salt Lake (Flowers), hard and soft hot, dry desert soils (Cameron, a single species, and phosphates 1959). Aquatic where light temperature 1961: Killian and frequently may grow prolifi-

forms,

cally in favorable seasons when nitrates cases release obnoxious toxins (Prescott, found 1953), in the and lower sublittoral springs where zone the in hot

are high and in some species have also been is low (Ruttner,

intensity

may reach 86 C. (Kaplan,

1956). Other aquatic habitats of metals and acids (Palmer, with wood, animals such recent algae that are the as sponges, on and able Eh in sewage,

can include industrial wastes with a high content 1959). More exotic habitats include associations corals, activity grow of range and snails. (Treub, and blue-green In barren, rocks, 1888), eroded soil, on that been light transmitting survive. and even in areas of lbund it has from

under to

comparatively blue-green determined

volcanic

it has been is

Furthermore, algae

-0.200to

+0.700 volts and the pH from 1.5 to 11 (Baas Becking et al., 1960). That they can resist desiccation for decades has been shown in the revival of species from old, stored latoroid which has desiccation Nostoc soils (Bristol, can develop remained has been 1919). Reproduction growth can be quite rapid, and oscilsoil lbrms macroscopic in a few hours on desert

dry for a number of years. Prolonged resistance to found in a dried herbarium specimen of nonspore-forming revived after 88 years of storage (Lipman, 1944),

commune

previously

and later revived press). Resistance combination even photosynthesize PROSPECTS Research ulation FOR

after an additional time period of 19 years (Cameron, in is also found to low temperatures. At-80 C., algae, in to survive, and at -30 C. to slowly (James, GENETIC 1955)."

with fungi as lichens have been found

ENGINEERING powerful tools for the manipnucleic

on bacteria

and their viruses has yielded of cells. Genes,

of the genetic

apparatus

composed

of deoxyribose

acid (DNA), determine determine the physical istic, for example,

the protein characteristics

enzymes of cells and these, to a large extent, of cells. Thus if a cell has a certain characterfrom ultraviolet radiation,

a capability

for rapid repair of damage

78

it is dueto the presencef a particulargene several o or genes. cellwhichlackssuch A a rapid repair mechanism would lack thesegenes. he collectionof physicaland T chemicalattributesof an organismis calledits phenotype,while the collectionof genesdeterminingan organism's phenotypeis called its genotype.The goal of genetic engineering the deliberatealteration of an organism's is phenotypeby changing genotype,either by causingsmallchanges a specificgeneor, more its in important,by integratingone or moreforeigngenes into the geneticapparatus of the organism. ature,of course,hasbeeninvolvedin geneticengineeringslongas N a DNA and evolution havecoexisted.Geneticengineering hasbeenusedfor many years by plant and animal breedersto produce strains having particular characteristics. The current interestin geneticengineering, however,reflectsthe recentdiscovery of powerful biologicaland chemicaltechniquesfor the creationof new genotypes. Thesetechniques includewaysof generating ewgenes n from pre-existing genes, nd for movinggenes a from donor to recipientorganisms, therebycreating newcombinations pre-existing enes. of g The primaryway that modifiedgenes canbegenerated from pre-existing enes g is by the process mutation in which small random changes the chemical of in structureof a genearebroughtaboutby exposure certainchemical r physical to o agents.Recentadvances understanding f genestructureandin methodsfor the in o
in vitro chemical de novo synthesis of genes, suggest extremely powerful, technique for generating new genes. an alternative, potentially

More important to the field of genetic engineering, and to the particular goal of the creation of an ideal oxygen-evolving organism for Mars, are techniques 6f moving about pre-existing genes among organisms, so as to create organisms to accomplish having a particuthis for millenia. lar phenotype. Nature has used sexual recombination

Other techniques have been developed in the laboratory. Transformation is a process in which DNA is extracted from donor cells and taken up by recipient cells. This foreign the progeny donor with DNA (containing apparatus the by the extremely of the desired trait. the genes recipient for desirable traits) endowing can be integrated these process vehicle cells. differing by which into from genes, into as a (e.g., gene is cells. genetic cell, thereby viruses into cells and their

Transduction

is a similar as the donor recipient

transformation incorporated potentially another, vehicle mentioned Morrow joined

use of certain effective, means

into the virus, can be moved organism it utilizes genes the By the into

Recently et al., 1973).

a new, and Referred techniques donor

of introducing

genes from one organism (Cohen from gene element, the from the a plasmid, ato,

quite dissimilar, engineering, introducing by etal., allowing 1974). for

has been described a specific cells. It differs specific complex

to as plasmid

small genetic of any

previous

use, in principle, use of certain

enzymes introduced

desired

to the plasmid

and the gene/plasmid 79

into recipient

Oncein the cell the gene/plasmidomplexcanforma stable,genetically c functional unit whichreplicates andis passedn to the progeny the originalrecipientcell. o of It is by judicious use of thesetechniquesthat an "ideal" oxygen-evolving Martianorganism mightbecreated. Until recentlyonly limited informationhasbeengathered the genetics on of blue-greenlgae, reflectionof both the difficulty in obtaining a a appropriate mutants, and in growingthe organisms the appropriate in media.Both theseobstacles have beenovercome andsomeinformationon the geneticsystems theseorganisms of is becomingavailableand, it may be assumed, ill continue to do so at an ever w increasing rate. Severaltechniques associated with geneticengineering avebeen h demonstrated a few species blue-greenlgae,.e.,chemical utagenesis, in of a i m sexual recombination,and transformation with DNA (Kumar, 1962; Bazin, 1968: Shestakov Khyen, 1970;Herdmanand Carr, 1971:Orkwiszewski Kaney, and and 1974;and StewartandSingh,1975).Anothertechnique, iral transduction, hile v w not yet demonstrated, quite likely (Padanand Shilo, 1973).The presence is of plasmids the blue-greenlgae in a hasnot beendemonstrated. The initial descriptionof sexualrecombination blue-green in algaehas been challenged: owever, everal ecentindependenteportsseemto haveplacedthis h s r r phenomenon firm ground.Chemically on extractedand purified DNA, aswell as DNA released cellsin growingculturesof blue-green by algae, asbeenusedfor h genetictransformation experiments. Transduction dependent is uponthe sensitivity of the cellsto a classof viruses called_'temperate" iruses. hese v T viruses havethe propertyof actingascarriers genes of between donorandrecipientcells.Temperate viruseshavebeenisolatedwhich can infect blue-green algalcells.The transferof genes betweenthe algalcellsby transduction, owever, asnot yet beenreported. h h The presentability to generate newgenomes the algaeby mutationandrecomin bination, coupledwith continued genetic research theseorganismsndthe related in a bacteria, uggests the designandconstruction organisms s that of specifically adapted to the variousMartiannichesare quite possible.The useof thesespecialstrains wouldbe critically importantin any attemptto generate anoxygenatmospheren o Mars.

80

9.

CONCLUSIONS

In Figure tile capacity considerations

1 a series of Mars presented

of questions in the body

was posed Answers of this

which to these

would

allow

an assessment generated

of

to be inhabited.

questions,

by the

report,

are outlined

below.

Clearly,
T

the prevent

lack

of

an from

oxygen being

atmosphere inhabited intense with by

by

itself

is The

enough diurnal
MARl IAN NO

to

Mars

man.

temperature and For dust Mars must be storms to be

fluctuations, can truly be dealt

ultraviolet by adequate

radiashield-

PRESENT

ENVIRONMENT SUPPORT LI HUMAN

tion, ing. factors

habitable, and

these

environmental atmosphere

moderated

a breathable

YES

generated.

The the
PRESENT MARTIAN _N_ NO ENVtRONMENT T _MLL SUPPORT ERRESTRIAL

lack

of oxygen

in the

atmosphere

will

not

prevent par-

growth

of all terrestrial must be considered

organisms: capable

anaerobic of existing

bacteria

ticularly Martian algae

in specific blue-green

L,EE' /
IYES

microhabitats. or lichens

Possibly, also

photosynthetic in specific

can survive

microhabitats.

While will
GROWTH IN A FOR HABIT PLANET HUMAN RESULT FIT NO

the modify

growth the

of

bacteria

is possible, so

such as to

growth allow

not

Martian

environment do not The would

human However, possible

habitation limited and they

because growth do

bacteria of

produce and rate

oxygen. lichens at which slow, is it for and

bhie-green oxygen.

algae

generate

YES

would example, This

accumulate 100,000 to is due

in the years

atmosphere to produce

be very

a breathable

atmosphere. effects the

a combination limitations. and 81 low

of environmental For temperatures example,

inherent ultraviolet

biological radiation

intense

are environmental

factorsthat will limit the extent andvigorof surface growth;andbiologically,the rateof lichenreproduction alsoveryslow. is
It is possible
CAN ORGANISM ENGINEERED GROW PLANET HUMAN & MAKE FIT FOR AN BE TO NO _1_

to envision far better

genetic organisms adapted

modification in such to the present

of specific a way that martian prothe

terrestrial they might

oxygen-generating be with Such the

environment
i I YES

regard

to survival, might an oxygen

growth, dramatically

and oxygen increase and

duction. probability decrease

modification time needed

of generating

atmosphere

greatly

for its generation. modifying conducive produce particularly Mars for human the Martian to an the oxygen seems climate extensive atmothe key

It is possible such
TO _TO BE MODIFIED SUPPORT HUMAN CAN LIFE" MARS AN_NO ,_

to envision be far

that

it might growth

more to of

biological sphere. to unlocking

necessary potential

Temperature the

manipulation

YES

habitation.

an

increase

Greenhouse in planetary surface changes of the

effects and temperatures. conducive 18 presents oxygen increase in

advective warming could bring about This increase could cause additional to the growth a suggested on the surthce sublime individual other years of terrestrial scenario Mars. A temperature and steps than might available allow water reliable for key to organisms. the producstep a level exist is a at in of or time

environmental

kind

Figure tion
1 2 BESt STRATEGY ECOLOGICAL CONSIDERATIONS 3 ISPECIFY TIME 4 OTHER /

of

an

atmosphere

moderate which the the

carbon liquid

dioxide

would for the

might

state. required

It is impossible

to give

estimates

times

in the scenario, that

indeed, spans changes reduce by man in either the time and the climate to periods. required storage shorter of

of the 10,000

scenario to or an

itself

to suggest be required. would Mars Mechanisms

100,000 oxygen oxygen

Radical for the

of Mars produce

the organisms would

considerably to be inhabited

atmosphere.

concentration in even

of atmospheric

On the 1. No terrestrial

basis has

of information been

currently

available to identified. reserves a more

the study the ability However, and the accurate

group of important

concluded Mars data are

that: not

fundamental, life

insuperable unequivocally the extent be acquired

limitation

to support

available, for example, caps. These data must bility of Mars 2. tory. forms, For but the

of water before

composition assessment

of the polar of the habita-

can be made. human time life to exist, might the creation of an oxygen by of tens present of thousands atmosphere of years. is manda-

Such

an atmosphere required

be generated

terrestrial

photosynthetic

is on the order 82

_o ....

t_

I u.l

T a. 0

Ii,
z

_o_

>-

o_
I,U 0 Z - _ I 5_

__i

I- I): 0 u. )l.u Im ix: f i 0 a. 0 m_ _Z_ _0

c _ ,

83

3. Climate

modification

leading

to an increase

in surface

temperature

might be by the need to to

carried out by a combination of advective and greenhouse effects initiated injection of polar cap sublimate into the atmosphere. This injection would be carried out for a long time at the expenditure of amounts of energy the amount of solar energy incident on Mars for several years. 4. Mechanisms of genetic engineering currently available could be used to construct organisms far better terrestrial organisms presently available. 5. Altering isms, or both, either would the Martian significantly environment decrease adapted

equivalent

or under development

to grow on Mars than those photosynthetic organsteps are

or available

the time required

to create an acceptable

human habitat on Mars. Indeed, it may be mandatory taken, Mars may well be made into a habitable planet.

to do so. If these

84

10.

APPENDIX LICHENS

A: AND

BIOLOGICAL CYANOPHYTES

CHARACTERISTICS ON MARS

OF

GROWTH

OF

GENERAL

MATHEMATICAL

APPROACH

To energy lichen of the -

determine balance has organism rate

the model

coupling representing

between two the the can with

organisms terrestrial assumption then

and

the

Martian

environment, algae temperature balance.

an and The to

organisms that

- blue-green the it is in energy as well

been

developed. is determined of the

Given

environmental

parameters,

transpiration photosynthesize. The solar rative source and local upon the and

organism

be determined

as its ability

major thermal

physical radiation, and

processes

controlling and

the

temperature energy atmosphere the organism with or

of

the

organism

are

convective thermal the solar solar 0.86

conductive from the varies

transport, provide depends

and evapoan energy upon equator depends and gas is the at season,

cooling. for the time noon the amount

Solar of the

radiation radiation and, ly/min. radiation The primarily absorb radiation this

organisms; Maximum is about

reaching

transparency

atmosphere

of course,

Martian occurs near

latitude, the radiation

of day. and

at the surface planetary that thermal 15/_m

thermal

atmospheric of CO2, flux by the a. incident gas which of about ability For on

temperature, which can absorbs 0.02 of the it, so the

of the lowest radiation. spectral

few kilometers, On region Mars and the

primarily an energy organism absorptivity, radiation

in the

produces to the by an as:

ly/min organism

at the surface. to absorb the absorbed this

Solar

radiation

is coupled which is given

radiation, absorbs the

all practical

purposes, radiation

organism by

all of the

planetary

organism

can be given

R =aQ

s +Qp

(A1)

where

Qs is the

incident

solar

radiation

and

Op the

absorbed

planetary

radiation.

85

Heat transporteither to or from the organism occursat the upperandlower surfaces thereis a difference temperature if in betweenthe organism andthe air or groundor both. At the uppersurfacemolecular onductiontransports c heatdown the temperature gradientacross laminarboundarylayerof thickness a cm which a 6
is related organism to a mean wind D tcm_ speed by the near the surface 2 V (cm/sec) and a characteristic dimension expression,

8a=

0.4(D/V)L/2

(A2)

If the then ism's diffused

near heat by

surface is conducted eddy

air temperature from the into than transport is lower energy the the

is lower organism near

than through

the the The

temperature boundary opposite

of the layer occurs

organism, and rapidly organfor this

atmosphere. surface Cto p is

if the

temperature

temperature.

An expression

convective-conductive

transport

Cto p = Kair(Twhere T, T a are the temperatures (K) of the

Ta)/6a organism and air: Kai r is the and with

(A3) thermal Kai r the

conductivity for COz, the major is approximately 35X 10 6 cal/sec Heat surface. transport by conduction analogous

constituent of the Martian cm deg at about 273 K. will occur to that above if the organism

atmosphere: is in contact

An expression

can be written,

Cbottom where mean model influences Two emission ing to its that is, A is a characteristic thermal and are conductivity. discussed the in the temperature allow cooling and solid half thickness next

= K(Tof the

Tg)t,_
organism are Molecular and K conduction

_A4) is a corresponding by the with biological the surface

These

parameters section. of the organism.

determined

strongly processes and

which

the

organism

to

rid

itself body

of energy radiates nearly

are energy

thermal accordbodies,

evaporative

(transpiration). and liquid

Every bodies

temperature,

radiate

as black

I = eoT 4

(A5)

2 Recent

work has shown

that the coefficient

is probably

closer to 3 than 0.4. However,

this

change does not significantly influence the cyanophyte results. In the lichen model, temperatures are more closely controlled by the surface, the lichen temperature being within 10 K of the surface. Water loss is larger but photosynthetic rates are about the same. 86

where unity; every

o is the Stephen-Boltzmann for example, gram of for water water evaporated

constant from

and e is the emissivity the emissivity about as, an organism,

which is about

is close to 0.95. For are

at room

temperature,

580 cal of heat

required.

This loss of heat energy

can be represented + ra)

E = 580 sdt/(rt

(A6)

where sdt is the saturation water vapor density (g/cm 3) at the temperature of the organism, r t is the internal resistance which the organism offers to water loss, and ra is the air resistance across the laminar boundary layer given by

ra = 6a/DH2 O-CO2 where DH2 O-CO2 is the diffusion is about 29.2 cm 2/see. The energy balance equation coefficient for H20 near the Martian symbolically, surface

(AT) and

can now be written

R = Ctop(T) and ature for a given biological of the organism model

+ Cbot(T)

+ I(T)

+ E(T) parameters,

(A8) the temperrate is given

and assumed

environmental

can be determined.

The water

loss or transpiration

by equation GENERAL

(A6), that is, E/580. ASPECTS OF MODEL variables, to be discussed subsequently,

With the exception

of several individual

the general mathematical formulae for both the cyanophyte the same. Treatments for both models are similar to those energy exchanges and photosynthetic (Gates, 1968; Gates, 1970; Nobel, energy exchange between the equation: R = eaT 4 + k(V/D) _/z (TTa) + K(TTg)/A

and lichen models are employed in studies of

production in leaves of higher terrestrial plants 1970). It is assumed that the complete budget of and its environment can be represented by

the organism

+ 580[sd t - (RH)(sda)]/(r

t + ra) (A9)

The

individual

variables

have

been

defined

elsewhere

(see pp. 85-87, terms. The

95-96). first

The term three

total energy surface, consists only

budget

for a hypothetical be summarized organism's

organism,

living on, or just below, The and emissivity.

the Martian

can therefore of the

as the sum of three

temperature 87

remaining

terms speed),

consider the

the last

energy and tcrm will

flux is of

restilting primary

from interest,

convection since to the

(air temperature respectively. involves variables cooling

and In the

wind this water which

conduction,

transpiration

tevaporative

coolingL it necessarily other the

discussion loss the

organisms the energy be

sustain

in response To

physical

characterize nent (r t) of the must

surface budget,

microclimate. the

evaluate resistance the total

evaporative of the resistance organism

compoloss to

characteristic Once

to water organism

measured

or

calculated.

of the

water flow is evaluated (the sum ofr t and the resistance, ra, due to the air layer above the organism), the flux of water vapor may be calculated first law of diffusion:

nonturbulent from Fick's

J = AC/R tt

(AIO)

where between R tt

= total the

water inner The

vapor portions model

flux, of can

AC = the the

change

in water and desiccation the

vapor

concentration and

organism the

atmosphere, stress a hypothetical

= r t + r a.

thus

estimate the of

organism known discussed

will be subjected or are calculated. for each case

to within Estimation

accuracy to the resistance

which these wthies for

parameters are both models is

subsequently. is the potential for photosynthesis that subject on rate (T), this of a terrestrial involves controlling although study CO2 group major organa comele-

Of critical ism on plex ments. more set the of The complete, the

importance surface.

Martian metabolic

It is acknowledged which are

photosynthesis to numerous higher model. intensity level, The (I),

interactions of

consideration is far beyond organism's include vapor which water

photosynthesis the scope photosynthetic temperature of the

perhaps chose limit-

present light

to represent ing variables, (Cco 2 ), and

as a function

of several

concentration (S). Further-

concentration,

or saturation, SI

of the organism

more, these "stibstrates" tions each proceed element

last three elements _1, CCO 2, and to enzymatically controlled reactions according is represented to Michaelis-Menton by a kinetic term

can be thought of (Gates, 1970). Since the individual

as limiting these reacinfluence of

kinetics, of the t'orm:

P=Pm

I(1

+ K/Xt

(A11)

where K = the S (I'(S)). organism, kinetic

photosynthetic

rate,

Pm=

maximum

possible

photosynthetic

rate,

"Michaelis Following as controlled terms

constant" for the assumption by these each

the reaction, and X = I,Cco 2, or a function of of Gates (1970) the photosynthetic rate of the three variables variable" is represented as the product of the

representing

limiting

88

P##l

P= (1 +K1/I)(I + K2/Cc02)(1 + K3/f(S))

(A12)

where K l, K2, and K3 are the constants applicable to I, CCO 2, and S, respectively. Before these parameters are further evaluated, the controlling infhience of the ten> perature (T) upon the process lntist first be considered. Temperature first is a specific can be thought effect; the of as influencing net photosynthesis rate in two ways. The is a function of maximum photosynthetic

temperature. Strictly speaking, this means that light compensation points change as a function of T also. A complex model would have to consider T as a variable in a set of differential equations used to describe respiration and photosynthesis. study group's Martian models, however, have an advantage over those which be formulated which mately 260-285 this temperature relatively low, for terrestrial conditions in this respect. The temperature the hypothetical organism may photosynthesize is limited The might

range over

to that of approxi-

K. Measurements of photosynthetic rates of terrestrial organisms in range show that photosynthesis and light compensation points are and they do not change drastically with change in temperature

(Lange, 1969; Lange and Kappen, 1972; Fogg and Than-Tun, 1960; Gates, 1968). Rates and light compensation points may change by less than a factor of about 5 (and usually by only reasons. 2 or 3 times). First, the This limited and range is not significant to the model from the for several accuracy the nature resistances, in their of magnitude constants, precision of the predictions

model depend entirely upon actual values for temperature, factors. tions The cumulative as small as one (resistances, error order

of the component variables, such as the thermal conductivities, and other physical makes it seem unlikely Second, the that variarates) are significant. and observed biological

estimation

parameters

"Michaelis"

photosynthetic

vary considerably among species, and therefore a mean value, derived for a hypothetical case, will inherently possess a large standard deviation. Finally, the ultimate organism(s) used for Martian atmospheric manipulation may have very different properties due to genetic engineering. These factors, combined with the fact that the experiments which were used for the estimation of the variables rarely approached "true" Martian conditions of temperature and pressure, justifies neglecting the variation of maximum "reasonable" only higher photosynthetic rates with temperature. The aim was to obtain contribute in (in some cases optimistic) values, and factors which would order effects (variations) to the models were not considered. generally provides a controlling influence

Temperature

on photosynthesis

the model. Photosynthetic production, as are all cellular metabolic processes, is sensitive to freezing. This factor, therefore, can determine the cutoff points in both cases. The physical property of solute induced 89 freezing point depression implies that

cytoplasm,which maycontainrelativelyhigh "solute" concentrations, remain will liquid (gel)at temperatureselow273K. In lichens,for example, et photosyntheb n sis has been observed low as 262K (Langeand Metzner,1965).The cutoff as temperatures the models for werechosen arbitrarilyto be at 260K and273K for lichensandalgae,espectively. r Giventhe general properties commonto both cases, a discussion f someof the individualfactorsfor eachmodelcantakeplace. o PARAMETERS OF
Figure organisms 8 shows CYANOPHYTE the proposed as existing MODEL schematic for the blue-green algal model. filaments. The They the in

are envisioned

in a thin layer of interdigitating

may be surrounded by a gelatinous covering and can reside at or just below surface of the planet. Since such algal mats are commonly crusty or "cuticose" their to the upper rapid segments, moisture a desiccated loss, which layer would is proposed to exist overlying result cells. This desiccated layer, it is suggested, may initially from exposure of the Martian

the "active" of cells environ-

be characteristic

ment. This layer may attain a thickness and this desiccated matrix might provide

equivalent to that of the active cell layer, an additional barrier to water loss by the

underlying cells. The resistance property of the desiccated cell layer is an important variable in the model. The thickness of the active cells has arbitrarily been chosen at 500/am. the This severe layer By terrestrial Martian is in intimate arbitrarily standards contact this is relatively conditions with the surface thin, however, permit it is assumed luxurient subsurface is proposed pressures that environmental would not growth. of Mars as being on Mars, it

or immediate The remainder

and consists water-saturated

of 50% cells by volume.

air. In view of the extremely

low atmospheric

was thought unreasonable to postulate a liquid water layer, would be of little consequence to this model. It was desired and resistivity upon (resistance certain per unit thickness) to establish property Resistances The water thetic of the other variables.

although its presence to vary the thickness the influence of this

to Water Transpiration diffusion and resistance thus indirectly is a critical affects that model the variable resistance since it directly temperature and influences photosynas the

flux rate.

organism's

Figure 8 indicates component

the overall in series.

can be represented

sum of several

resistances

This representation

is similar to that

employed by Nobel (1970) in his discussion of leaf transpiration. The total resistance to water vapor diffusion fRtt_ is equivalent to the sum of the resistances of the cell wall covering layer _Rdcl_ and that the organism
(Rwv).
c/

(Rwv), the intercellular air space "-wv"_Rias_ desiccated the cell due to the undisturbed boundary air layer immediately above
Values

CW

are estimated 90

for these individual

resistances

as follows.

All water the cell wall atmosphere

vapor covering,

diffusion the through

occurs the

upwards air space

from and

the

saturated layer).

cell walls, layer, to the

through outside

intercellular

desiccated

(crossing

nonturbulent + Rias "'wv path diffusion from path

air boundary + Rdcl "wv the + Rw_, a cells

RtOt _ Row wl' "'wv The space layer is 6 a. RCWwv is dependent gelatinous incorporated resistance
CW

(AI3) the the intercellular nonturbulent air air

effective the

mean effective

diffusion mean

through through

is 6ias;

length

upon cell 0.2 wall

the the

composition cells. In 1970). mesophytes, degree (Nobel, for a high

of plant

any

layer a this up

(e.g., waxy

cuticle, layer to

sheath may vary be in

matrix) into from 0.1

surrounding the to

leaves,

Since and

is reported to 2 sec/cm

sec/cm

for

xero-

phytes,.Rwv = 2 sec/cm (assuming RlaS can be calculated wv from

of drought

adaptation).

Rias = 6ias/DHz wv

(A14)

where OH20 is the diffusion coefficient for water vapor in Martian air. Assuming _iias is 50Y(_ of the thickness of this layer, then 6ias = 250/am, Rwr.as _- ,.50/"_DH20. i R dcl is designated as a variable resistance, since its value is a model parameter. "" W !' It may numbers layer be dependent upon of cells dying, and matrix (desiccated adaptation that the cells of the a number the precise are cell of properties; composition to be for example, age of the of the dead (or desiccated) physiologically sheaths) inactive). mat, cell a

assumed wall (and/or

Since

xerophytic logical

surrounding

is assumed, large:

it is

resistance

to water

vapor

diffusion

be significantly

perhaps

equivalent to the cutinized layer in xerophytic plant leaves, the 200sec/cm range. Therefore a maximum resistance maximum sented layer as the thickness product of 500/am resistivity were selected. layer This _dcl, .rW],
_

which have resistances in of 1000sec/cm and a resistance and and can be repre(ldc/_ from 0 to

of the where

of the from 200

its thickness 1dcl varies

"wrRdCl_" (r d_i( )( 1dcl), = 500pro. The speed, and diffusion

_dcl varies rwv

to _ E+4, ,.

pathway

through from this

the

nonturbulent Thus, 20

air layer

(6 a) varies using"

with

wind

can be calculated

value.

Ra_,l, is calculated

Ra wl' = 6a/DH The explained total above, resistance to the diffusion as:

(Al4a) vapor for the cyanophyte mat, as

of water

can be summarized

RtOt_ WI_

RWF

cw + _'Wp Rias

+ Rdcl "'Wp

Rwv

cw = Rwv + [(DH20)91

_ (6ias

+ 6a)]

+ , dcl lrwv

idcl)

(AI5)

Cyanophyte Mat Resistances COz to

The area the the ratio mat of the surface mat) area area of the increase fact that (Acells/A can be

Diffusion cells within the mat to the exposed surface for from lies diffuof the by this

of the

used

as a correction for CO2

factor diffusion.

to compensate This results

of surface the ultimate inside must also

which

is available for CO2 Thus the CO2, cell

"destination" the cells.

(a substrate in addition

for photosynthesis) to the water vapor

at the sion cell.

chlorophyll pathway, This necessitates path and the

diffuse

past

wall and

through resistances saturated

the cytoplasm presented cell of CO2 wall,

the

estimation such total

of tile additional structures resistance by water as the

increased lemma, the

length cytoplasm.

through The

plasmaof

to the diffusion vapor + a RCO2

is the sum

following: a. resistances also encountered cw (Rco2 b. c. d. cell wall resistance plasmalenmla cytoplasmic

+ oias _'C02

+ odcl "'CO2

(RU_o2) (R_to) 2 (R_002) similar to Nobel's A mat ..... (1970) analysis:

resistance resistance

Proceeding

in a fashion

2kx.j . (A16)

where

R/CO 2 _,,.i

resistance

of the/th

barrier

to CO 2 diffusion

thickness diffusion

of the/th constant

barrier of CO2 in the/th barrier

partition

coefficient

for

the barrier 1.0 (Nobel, 1970), and Ar w

Assuming (thickness

Dw _-- 5 E-6 cm 2/see, C02 of the cell wall) is 0.1 _m, Rw CO2

and

/_CO 2"_'*'

_ 0.01

sec/cm

(ifAmat/Acells

= 1/200). 92

Plasmalemma

Resistance

to CO2 Diffusion (1970) analysis: assuming that the

In a fashion permeability cells, which Cytoplasmic Using

which is again similar to Nobel's

coefficient for,. is at least Resistance (A16) where

CO2 entering algal cells &COo ) _ CO2 entering plant 0.01 cm/sec, R/O2,_ -- (Amat/Acells)/Pco 2 = 0.5 sec/cm. to C02 Diffusion

cp cp = A_ cp = 1 E-5, DCO 2 = 1 E-5 cm2/sec, KCO z 1, and cp Amat/Acell = 5 E-3, then RCO 2 _ 5 E-3 sec/cm. It is therelbre concluded that the additional resistances to CO2 diffusion (R w pl CO2 + R,_,,,
2

+ R c 2) are negligible ,_,_,7"Pc_

compared

to the magnitudes

of _DCW 2 + I,/\ CO

"CO2tas+ R_ 2 + R_O 2). It is assumed that the resistances to diffusion of water vapor and COz are inversely related to the ratio of their diffusion constants (Nobel, 1970). At Martian surface pressures the diffusion constant of COz in CO2 (Dco 2 ) is

22.3 cm2/sec, and the constant for water resistance of the jth barrier to CO: diffusion water vapor diffusion by the equation:

in CO2 (DH2 O) is 29.2 cm2/sec. The can be calculated from its resistance to

R_O 2 = Rwv(DH2 ] The total resistance ptot

*'CO 2

o/Dco2

J _CO 2 ) = 1.3 1 Rwv as follows:

(A 17)

to CO2 diffusion + rfias "'CO2 + 1adcl "'CO2

can be represented

_
=

cw RCO2

+ Ra 1.3 cw ias dcl a CO2 = I(Rwv + Rwv + Rwv + Rwv)

Estimation The uptake)

of Substrate equation

Constants

for Photosynthesis rate (equivalent to the net rate of CO2

for net

photosynthetic by equation

has been given previously

(A12).

The third term of the denomi-

nator of this equation applies only to the subsequent lichen model; therefore, K3 = 0 for the cyanophyte model. To complete the model, three parameters (P, K1, and K2) must be estimated. experimental experimental conditions, Unfortunately, these parameters must be evaluated from data contained in the physiological literature on blue-green algae. The conditions used to derive these data did not reflect Martian surface so that the results derived from using these reasonable indicative values must be interpreted values, and not to Mars colonization or plausible of an actual

with caution. It was desired to obtain achieve results which would be truly

93

experiment. estimates

The

same

quantitative parameters

approach become

can available.

be employed

when

more

accurate

of tile critical

Using the data of Kratz and Myers (1955), Pnz was estimated from a manometric experiment to be 8 E-7 moles (CO2 or O2)/cm 2 surface coverage/hr. This relies sents gave CO2 values K2 tions their are upon 800 plots the estimate or the of a mat study which rate were are the 4 E-6 taken molar, used group that 1 g dry cm thick. wt of algal same culture repre-

cm 2 coverage of net

is 0.05 light estimate

These and

investigators versus If the of K1 and the predic-

photosynthetic which constants at one-half

versus to

intensity KI and

photosynthesis respectively. to the the vahies out that

concentration, of these 6/:'-3

K2,

to be numerically photosynthetic respectively.

equivalent rates, It is pointed

"substrate"

concentrations

maximum

ly/min

and

of this model are sensitive estimation were derived for only several

to these values from laboratory species (at 298

for Pm, K 1, and K2. The data used in cultures incubated under "optimaF' K). In a sense, these the may represent

conditions unrealistic sensitive originally these net errors

values (i.e., maximal) for Mars. On the other hand, to the (1 g dry wt _ 800 cm 2) surface area assumption, derived may on a per interact production needed gram either dry which weight basis. It should by the energy of water point from cells has antagonistically is predicted the

actual P#n value is since the rates were that all of the the to influence This and concludes photosynthetic which the The will by cells model be sus-

be recognized model.

or synergistically

photosynthetic

estimation production Since can sustain assumes tained tion. At

of all variables

to calculate model. the extent cut-off

budget loss been has

for the cyanophyte mat it is difficult to estimate photosynthesis, the point The 105; of the the total cells

tinder

an arbitrary rate, water are assumed

selected. (AI2), lost state

that until this

photosynthetic

calculated

equation been

content

of the

transpira-

to be in a quasi-dormant to remain sunrise. day the life, At this and

of metabolic throughout the photowater The cells

shut-down. the cells remainder can

"active" of the the day, water normal for

cell layer until lost rate. the the

is hypothesized following the

in this state time,

prestimably, with

regain at their

during

previous

can proceed it, on 273 of

synthesis source" can also the dent

Parenthetically, of any

presence

of a "subsurface Mars.

is required be shut-down change The of the of

existence

as we know of less than model the during by can rate which the

by an organism in radiation diurnal flux lluctuations The

temperature on Mars, will period be can the control of time computed then

K. By estimating a time depenthe loss and

diurnal feature.

be given water they

temperature cally capable

organisms.

are physiologiThe the net prod-

photosynthesis production

can per day

above as the time as the

criteria. sum of

photosynthetic ucts of the

be calculated over short

instantaneous This sohition zero.

photosynthetic approaches

rates a true net

intervals limit

of active of the time

metabolism. intervals

production

approaches

94

The authors

acknowledge

the fact that this approach

is appropriate

in view of

the large errors which are inherent in the estimation of some of the critical parameters. They stress that this type of model can be improved immensely in order to make Mars, research standpoints. elled. K1, Second, K2, K3, it appropriate which on the First, to more accurate from currently of the metabolic of the critical for those biological there aspects is a need determinations planned model of the physical probes. They urge from be conducted parameters that at least description of two of will result further

for a more

rigorous activities organisms

mathematical

the photosynthetic etc.)

and respiratory evaluation should be made

of the organisms parameters which

to be mod(e.g., Pro,

a better

photosynthetic

may be considered

primary candidates for Martian inoculation. Experiments face conditions should be run to obtain better estimates models. tions The coordination to all future OF of these planetary two research engineering efforts pertinent

under simulated Mars surof these variables for future will provide future predicand design studies.

feasibility

PARAMETERS

LICHEN representation

MODEL of the study group's lichen model is presented in

The schematic

figure 7. Its structure and the representation of the variables are very similar to those employed in the algal model already discussed. As the diagram indicates, the lichen is envisioned as growing on the Martian surface, and is fully exposed to solar radiation. It could also be buried just beneath the surface, covered with sand. The segment upper (25% including cells surface. presents Inputs model. except active of the surface, the lichen which is modelled is a rectangular symbionts parallelepiped of this segment in a homogeneous layer over of square is Tcm, matrix the top with an edge length layer of active and a very of 1 cm. The thickness thin "cuticose"

algal and fungal

by volume),

protective

Its total thickness can vary from 0.05 to 0.3 cm. A nonturbulent air layer an additional boundary to diffusion, and is a function of the wind speed. to and The that outputs from the system are the at which same as those the organism in the cyanophyte previous case, basic general assumptions temperature are also the same as in the temperature 1972).

the minimum

is photosynthetically is reasonable

is set at 260 K (as opposed observations

to 273 K). This cut-off (e.g., Lange and Kappen,

in light of published Resistance Rather algal model, (1927)

to Water Vapor

Diffusion as was done for the empirically. Stocker genera, Usnea and

than calculate individual resistance components, the total effective resistance (R l) was estimated water loss and uptake rates for two lichen

reported

95

Parmelia.

These

data vapor is:

were flux.

used The

to estimate rate (E)

a reasonable water

value vapor

for

the

overall from

resisa plant

tance to water leaf by diffusion

at which

escapes

E =" where

_d t - RH(s.d a i ..... _. i r t + ra

(A18)

sdt

water

vapor

density

within

the

organism

considered

saturated

at the

organism RH

temperature

relative

humidity

of the

surrounding

atmosphere

saturation

vapor

density

of the

surrounding

air at the air

temperature resistance of the organism to water vapor diffusion

rt

ra
Solving data face water to two diffusion area

the

diffusion

resistance

of the

external

pathway

this

equation (1927) This

for

the

total lichen under the

resistance segment the

(R tt yields

= r t + ra) a total that

and

applying

it to the vapor to area span one

of Stocker of 4.6 covered. vapor orders

for the seems but factor.

resistance 1 gm dry value (dry

to water

E-2 min/cm,

assumption is sensitive to

wt = 25 cm 2 surresistance

to be a reasonable value

empirical the

for the

diffusion,

weight):(surface factor may

coverage)

conversion

Unfortunately,

the variation

in this

The wind

of magnitude (Blum, 1973). l The estimate of Rwv, also allows the calculation of R/O2_ by equation (A17). resistance (R a) due to the nonturbutent air layer can be calculated from the speed as noted for critical Lichen previously. Photosynthesis for equation for the the (A12) algal can be estimated The major saturation (Lange, gives 1.25 same for from factors of data which the 1969, some and which contained influence thalli, and Desert moles/ given

Parameters The in the the

variables

literature,

as was done rate are

model.

photosynthetic CO2 1972). species,

percentage and which light Lange

water

lichen Lange Negev 1.4E-6 it was

temperature, Kappen, lichen cm 2/hr, previously

concentration, the Pm data were parameter

intensity (1969) to be the

From KI and

estimated K2

ly/min value

respectively. (4 E-6 M).

The

retains

96

The final parameters hich must be evaluated w are

saturation synthesis, followed similarly tion were in equation as a function first to the order way modelled. uses this content (S) dry I gm and (AI2). of The the data of" Lange of water kinetics. in the kinetic by the amount (1969) saturation This degree zero the last order influences term

those showed of

pertaining that the has and of The lichen thallus, been CO2

to water photoroughly

then The

effect intensity

represented concentrainfluwater as a

in which

of light

denominator approach. K3

equation(Al2), controlling f(S). lichen weight middle as the water point The thallus

[ 1 + K3/f(S)], ence of water

"Michaelis-Menton" is represented is defined as the example, Since S= range, complication Since the the lichen content, cm thick

parameters of water

and

saturation percentage thallus observed possible to permit occur

variable of contains maximum water active its

in the

weight. of

For

S = 100fTc if 1 gm dry 100% this falls within selected minimum cut-off must water f(S) 80f_ the value is that expression segment, was the

of lichen of the inaximum saturation does not to

water. saturation

water metabolism

content. percentage for this. (Kappen,

A further water

is 20%.

photosynthetic

at zero

be formulated loss (the assumed

compensate cut-off covered. point)

In a 0.3 1973)

occurs

at a net

loss of 40 mg water/era

2 surface

area

Therefore, f(S) 40(mg ....................... H20 40 lost per cm 2) X 100 for S>20 (A19)

This

concludes From the

the estimation standpoint pertaining be conducted parameters. of to

of all important inherent the which error,

parameters the model. study

in the group more

lichen offers reliable

model. the that same cauof engitinder

tions

as those

cyanophyte are designed important, done

It is suggested

laboratory estimates

investigations the critical

to obtain respect the

It is also work be

with

to Martian viability of

planetary lichens

neering, anaerobic

that

further

concerning

conditions.

97

11.

APPENDIX

B:

GREENHOUSE

EFFECT

The carbon chapter A uniformly sphere cm 2 sec. the above

mean dioxide 5: this mean

global and

change water

in surface in the the

temperature atmosphere

which of used Mars

would has

be affected been discussed

by in

vapor

appendix thermal

describes profile until absorbed that for all

methodology (Noll

for those McEIroy, flux at the

calculations. 1974) top as and of the shifted atmos ergs/ so that true, by

is assumed the solar carbon the outgoing

and

in temperature balances value for the It is assumed is fixed vapor

planetary which dioxide The absorbs

radiation, dioxide carbon effect. absorptivities 1967). For

is taken calculations.

1.2810 radiation This

negligible

solar

is not radiation

however, water ing amount 1.3810 The equation,

a water was to 10

greenhouse from the

absorption for the eight

of solar water increases

vapor from 0.8 of

calculated 6.3/_m mbar, 2 sec. flux

bands water to

extendvapor about the

(Rodgers, the at the

the

assumed radiation

maximum

absorbed top of the

solar atmosphere,

s ergs/cm planetary

F _', was determined

from

where or eight Planck tion

the

mass interval

path vapor) surface flux

is referenced spectral to the

from intervals and mass

the the used

surface; index

T/is

the

flux

transmissivity and Boa(u) A is the

for are the by the

a spectral

of width

6 i, where

i runs

over

the Bed(o)

six (carbon and

dioxide) contribu-

(water functions the

in this study; u, respectively; undergoes body which

for the surface

path

from

outward

flux

no attenuation The and

atmosphere;

this

can be represented

by black

radiation.

wave-number the integration are from

integration was over transmission The McClatchey spectral

performed by a six point Gaussian was the sum of finite differences. data required The to carbon evaluate dioxide 99 the band

quadrature, transmission which

functions

et al. (1973).

is of primary

importance

occursnear 15/xmwith spectral idth of 500 to w

tion tar consists infrared of two region bands, extending and model path intervals one at 6.3/_m out the and from lines t1250 about in each profile

862 13/xm.

cm -_ . The Each grouped used. was give

water

vapor the other

absorpin the into

to 2450

cm -_ ) and band into Also good

was divided five

several decades. line account. laboratory

spectral on

intervals, band the

were were

line-strength of into with taken

A random The

Voigt

the dependence agreements

strength

mass

weighted were chosen

temperature so as to

explicitly

spectral

measurements. atmosphere temperature the mixing was for ratio divided each into layer sixteen layers of equal to be that for each pressure for the calculation. increments. center of the

The The layer, mean and

was assumed to be constant

was assumed

1O0

12.

REFERENCES

Ahmadjian, V., and Hale, M. E., eds. (1973). The lichens. Academic Press, New York. Alexander, M. (1961). Introduction to Soil Microbiology, John Wiley & Sons, New York. Barth, C. A. (1974). The Atmosphere of Mars. Ann. Rev. Earth and Planet. Sci. 2:333 367. Bates, D. R., ed. (1964). The Planet Earth. Pergamon, New York, 175-177. Bazin, M. J. (1968). Sexuality in a blue-green alga: Genetic Recombination in Anacystis mMulans. Nature 218:282-283. Becquerel, P. (1948). vide et deux 226:1413-1415. Reviviscence du Xanthoria parietina dess_ch_ avec sa faune, six ans dans le semaines a - 189C Ses consequences biologiques. C. R. Acad. Sci. concentration in the

Berkner, L. V., and Marshall, L. E. (1965). On the origin and rise of oxygen Earth's atmosphere. J. Atmos. Sci. 22:225-261. Bliss, L. C., and Hadley, Bot. 51:870-874. Blum, E. B. (1964). Photosynthesis

and respiration of Alpine Lichens. Amer..I. V., and Hale, M. E. (eds.) In: Microbial Growth. 19th

O. B. (1973). Water relations. In: The Lichens. Admadjian, Academic Press, New York, 381 400. Brock, T. D. (1969). Microbial Growth Under Extreme Conditions.

Symposium of the Society for General Microbiology, Cambridge University Press, 15 41. Butin, H. (1954). Physiologisch-6kologische Untersuchangen (iber den Wasserhaushalt und die Photosynthese bei Flechten. Biol. Zentralbl. 73:459-502. Brock, T. D. (1973). Evolutionary and Ecological Aspects of the Cyanophytes. the blue-green algae. Carr, N. G., and Whitton, B. A. (eds.). University Berkeley, 487-500. zur Okologie and Physiologie 160:Abt. B:72-99. of Representative In: The Biology of of California Press, bei einegin Sci.

Btittner, R. (1971). Untersuchungen Strauchflechten. Flora (Jena) Cameron, R. E. (1963). 108:412-420. Cloud, Cohen, Morphology

des Gasstoffwechsels

Blue-Green

Algae. Ann. N.Y. Acad. Proc. Nat'l.

P. E. (1965). Symposium Sci. USA. 53:1169-1226.

on the evolution

of the Earth's

atmosphere.

Acad.

S. N., Chang, A. C. Y., Boyer, H., and Helling, R. (1973). Construction of Biologically Functional Bacterial Plasmids in vitro. Proc. Nat. Acad. Sci. USA. 70:3240-3244.

Conrath, B., Curran, R., Hanel, R., Kunde, V., Maguire, W., Pearl, J., Pirraglia, J., and Welker, J. (1973). Atmospheric and surface properties of Mars obtained by Infrared Spectroscopy on Mariner 9. J. Geophys. Res. 78:4267 4278. Crutcher, H. L. (1969). Temperature and Humidity in the Troposphere. hi World Survey of Climatology, Vol. 4: Climate of the Free Atmosphere. Rex, D.F. (editor), Elsevier, Amsterdam, 45-83. 101

Fanale, P.,andCannon, A. (1974). xchange absorbed F. W. E of HzOandCO2 between the regolith atmosphereMars and of caused changessurface by in insolation. Geophys. Res. J.
79:3397-3402. Fanale, F. P. (1975). Regolith storage of volatiles and an Earth-analogous Mars degassing model. EOS, Trans. Amer. Geophys. Union. 56:406. Foster, T. L. (1973). Response of selected microorganisms to a simulated Martian environment. Dissertation for Doctor of Philosophy, Texas A&M University. Fogg, G. E., Stewart, W. D. P., Fay, P. and Walsby, A. E., editors (1973). The blue-green algae. Academic Press, New York, 317. Fogg, G. E. and Than-Tun. (1960). Interrelations of photosynthesis and assimilation of elementary nitrogen in a blue-algae. Proc. Roy. Soc. B. 153:111 -127. Gates, D. M. (1970). Synergisms of life and climate. In: A Century oJ Weather Progress, edited by James E. Caskey, Jr., American Meteorological Society, Washington, D.C., 120-129. Gates, D. M. (1968). Toward understanding ecosystems. Advances in Ecological Res. 5:1-35. Gierasch, P. J., and Toon, O. B. (1973). Atmospheric pressure variation and the climate of Mars. J. Atmos. Sci. 30:1502-1508. Gierasch, P. J., and Goody, R. (1968). A study of the thermal and dynamical structure of the Martian lower atmosphere. Planet. Space Sci. 16:615--646. Gifford, F., Jr. (1956). The surface-temperature climate of Mars. Astrophysical J. 123:154 161. Godnev, T. N., Khodasevich, E. V., and Arnautova, A. I. (1966). Biosynthesis of pigments in lichens and overwintering plants at subzero temperatures. Dokl. Akad. ,Vauk. 167:451. Akad. Nauk SSSR Biochemistry 167:90. Godward, M. B. E. (1962). Invisible Radiations. In: Physiology and Biochemisto' of Algae. Lewin, Goody, R. A. (ed), Academic Press, New York, 551 566. R. M., and Walker, J. C. G. (1972). Atmospheres. Prentice-Hall, Inc., Englewood Cliffs, New Jersey. Hand, R. A., Conrath, B. J., Hovis, W. A., Kunde, V. G., Lowman, P. D., Pearl, J. C. Prabhakara, C., and Schlachman, B. (1972). Infrared spectroscopy experiment on the Mariner 9. Mission: Preliminary results. &'ience 175:305-308. Herdman, M., and Cart, N. G. (1971). Recombination in Anao, stis nidulans mediated by an extracellular DNA/RNA complex. J. Gen. Microbiol. 68: XIV. Holm-Hansen, O. (1963). Algae: Nitrogen fixation by antarctic species. Science 139:1059-1060. Horowitz, N. H., Cameron, R. E., and Hubbard, J. S. (1972). Microbiology of the dry valleys of antarctica. Science 176:242-245. Ingersoll, A. P. (1974). Mars: The case against permanent CO2 frost caps. 3.. Geophysical Res. 79:3403-3410. Jahns, H. M. (1973). Anatomy, morphology, and development. In: The lichens, Ahmadjian, V. and Hale, M. E. (eds.), Academic Press, New York, 3-58. James, P. F. (1955). The Limits of Life. J. of the British h_terplanetarv Soc. 14:265-266. Kappen, L. (1973). Response to extreme environments. In: The Lichens. Ahmadjian, V., and ltale, M. E. (eds.), Academic Press, New York, 311 380. Kershaw, K. A., and Rouse, W. R. (1971). Studies on lichen-dominated systems. I. The water relations of Cladomia alpestris in spruce-lichen woodland in Northern Ontario. Can. J. Bot. 49:1389-1399.

102

Kieffer, . H.,Chase, H S.C.,Jr.,Miner, .,Mtinch, E G.,andNeugebauer, G.(1973). Preliminary reporton infrared radiometric measurements fromtheMariner spacecraft.Geo. Res. 9 J.
78:4291-4312. Kliore, A. J., Fjeldbo, G. F., Seidel, B. L., Sykes, M. J., and Woiceshyn, occultation measurements of the atmosphere and topography Extended mission 78:4331-4351. coverage of polar and intermediate P. M. (1973). S Band radio of Mars with Mariner 9: J. Geophys. Res.

latitudes,

Kratz, W. A., and Myers, J. (1955). Photosynthesis and respiration of three blue-green algae. Plant Physiol. 30:275-280. Kumar, H. D. (1962). Apparent genetic recombination in a blue-green alga. Nature Dec. 15, 196:1121-1122. Lange, O. L. (1969). Experimentell-iskologische Untersuchungen an Flechten des Negev-WiJste. I. CO2-Gaswechsel von Ramalina macriformis (Del.) Bory unter kontrollierten Bedingungen im Laboratorium. Flora (Jena), Abt. B: 158:324-359. Lange, O. L., Schulze, E.-D., and Koch, W. (1970). Experimentell-iSkologische untersuchungen an Flechten der Negev Wtiste. II. CO2-gaswechsel und Wasserhaushalt von Ramalina macriformis (Del.) Bory am naturlichen Standort w_ihrend der Sommerlichen Trockenperiode. Flora 159:Abt. B:38-62. O. L., and Kappen, L. (1972). Research Series 20:83-95. Photosynthesis of lichens from Antarctica. Antarctica

Lange,

Lange, O. L. and Metzner, H. (1965). Lichtabhangiger Kohlenstoff - Einbau in Flechten bei tiefen Temperaturen. Naturwissenschaften 52:191. Leighton, R. B., and Murray, B. C. (1966). Behavior of carbon dioxide and other volatiles on Mars. Science 153:136-144. Leovy, C., and Mintz, Y. (1969). Numerical of Mars. J. Atmos. Sci. 26:1167-1190. simulation of the atmospheric circulation and climate

Lettau, H., and Lettau, K. (1969). Shortwave radiation climatonomy. Tellus 21:208-222. Levine, J. (1975). Argon on Mars!: Where is the water? EOS. Trans. Amer. Geophys. Union 56:405. Levine, J. S., Kraemer, D. R., and Kuhn, W. R. (1974). The distribution of solar radiation incident on Mars and the Jovian Planets. EOS. Trans. Amer. Geophys. Union 55:343. Lund, J. W. G. (1962). Soil Algae. In: Physiology and Biochemistry of Algae. Lewin, R. A. (ed.) Academic Press, New York, 759-770. McCormac, B. M. (editor) (1971). Introduction to the Scientific Study of Atmospheric Pollution. D. Reidel Publishing Company, Dordrecht-Holland. McClatchey, R. A., Benedict, W. S., Clough, S. A., Burch, D. E., Calfee, R. F., Fox, K., Rothman, L. S., and Gating, J. S. (1973). AFCRL Atmospheric Absorption Line Parameters Compilation. AFCRL, L. G. Hanscom Field, Bedford, Mass. AFCRL-TR-73-O096. Morita, R. Y. (1975). Psychrophilic bacteria. Bact. Rev. 39:144-167. Morrow, J. F., Cohen, S. N., Chang, A. C. Y., Boyer, H. W., Goodman, H. M., and Helling, R. B. (1974). Replication and transcription of eukaryotic DNA in Escherichia coli. Proc. Nat. Acad. Sci. USA 71:1743-1747. Nawrocki, P. J., and Papa, R. (1963). New Jersey. Nobel, P. S. (1970). In: hltroduction Francisco, 314-338. Atmospheric to biophysical Processes. Prentice Hall, Inc., Englewood Wh. H. Freeman Cliffs,

plant physiology.

& Co., San

103

Noll,R.B.,andMcElroy, M.B.(1974). Models ofMars' atmosphere. Space NASA Vehicle esign D Criteria (Environment), SP-8010. NASA Orkwiszewski,G.,andKaney, R.(1974). enetic K. A. G transformationtheblue-green of bacterium Anaeystis nidulans. Arch. Microbiol. 98:31 -37.
Padan, E. and Shilo, 37: 343-370. M. (1973). Cyanophages viruses attacking blue-green algae. Bact. Rev., Pintner, I. J., and Provasoli, L. (1958). Artificial cultivation of a red-pigmented alga, Phormidium p('rsicinum. J. Gen. Microbiol. 18:190-197. Pollack, J. B. (1975). "Mars." In: Sci. Amer. Sept. 233:106 117. marine blue-green

Quispel, A. (1959). Lichens. In: Handbook der pflanzenphvsioh_gie, W. Ruhland (ed.), SpringerVerlag, New York, 11:577 604. Rao, D. N., and LeBlanc, F. (1966). Effects of sulfur dioxide on the lichen alga, with special reference to chlorophyll. Brvologist 69:69-75. Rasool, S. I., Hogan, J. S., Stewart, R. W., and Russell, L. H. (1970). Temperature distributions in the lower atmosphere of Mars from Mariner 6 and 7 radio occultation data. J. Atmos. ScL 27:841- 843. Rasool, S. I. (1971). Planetary Holland, Amsterdam. 369 Reid, A. (1960). Stoffwechsel atmospheres. 399. und und In: Exobiology, C. Ponnamperuma (ed.), North

Verbreitungsgrenzen

yon Flechten.

II. Wasser- und Assimilabenachbarter

tionshaushalt, EntquellungsStandorte. Flora 149:345-385.

Submersionsresistenz

yon Krustenflechten

Richardson, D. H. S. (1973). Photosynthesis and Ahmadjian, V., and Hale, M. E. (eds.), Academic Rodgers, C. (1967). The Radiative Heat Budget of Massachusetts hrstitute of Technology, Dept. of Report no. A2, Oct. 1, 1967.

carbohydrate movement. In: The lichens. Press, New York, 249-288. the Troposphere and Lower Stratosphere. Meteorology, Planetary Circulations Project

Sagan, C. (1971). The long winter model of Martian biology: a speculation. Icarus 15:511-514. Sagan, C., and Pollack, J. B. (1974). Differential transmission of sunlight on Mars: biological implications, h'arus 21:490- 495. Sagan, C., Toon, O. 181:10451049. B., and Gierasch, P. J. (1973). Climatic change on Mars. Science

Schopf, J. W. (1974). 5:119-135.

The development

and diversification

of Precambrian

life. Origins of Life

Sellers, W. D. (1965). Physical Climatology. Univ. of Chicago Press, Chicago, Ill. Shestakov, S. V., and Khyen, N. T. (1970). Evidence for genetic transformation Ana_),stis nidulans. Molec. Gen. Genet. 107:372-375. Siegel,

in blue-green

alga

S. M., and Daly, O. (1968). Responses of Cladonia rangiferina to experimental stress factors. Bot. Gaz. (Chicago) 129:339-345. Singh, H. N., and Shrivastava, B. S. (1968). Studies on morphogenesis in a blue-green alga. I. Effect of inorganic nitrogen sources on developmental morphology of Anabaena doliolum. Can. J. MicrobioL 14:1341 - 1346. Sinton, W. M., and Strong, 131:459-469. J. (1960). Radiometric Observations of Mars. Astrophysical Journal on

Stewart, W. D. P., and Pearson, H. W. (1970). Effects of aerobic .'rod anaerobic conditions growth and metabolism of blue-green algae. Proc. Roy. Soc. kond. B. 175:293-311.

104

Stewart, W.D. P.,andSingh, H.N.(1975). Transfer nitrogen-fixing of (NIF)genes thebluein greenlgaostoc muscorum. Biochem. Biophys. Res. Comm. 62:62-69. a N
Stocker, O. (1927). Physiologishe und _Skologische Untersuchungen an laubund Strauchflechten; ein beitrag zur experimentellen 0kologie und Geographie der Flechten. Flora 121:334-415. Stone, Syers, P. H. (1972). A simplified radiative-dynamical atmospheres. J. Atmos. SeL 29:405-418. model for the static stability of rotating

J. K., and lskandar, I. K. (1973). Pedogenetic Significance of Lichens. Ahmadjian, V. and Hale, M. E. (eds.), Academic Press, New York, 225-248. In: Antarctic Ecology.

In: The Lichens, M. W. Holdgate

Ugolini, F. C. (1970). Antarctic soils and their ecology. (ed.), Academic Press, New York, 2:673-692. Vallentyne, R. R. (1963). 108:342-352. Van Baalen, C. (1961). 133:1922-1923. Environmental Vitamin Bl2 biophysics

and microbial of

ubiquity.

Ann. N.Y. Acad. Sci. alga. Science algae in of the of rice of

requirement E. (1971). S. (1967).

a marine

blue-green

Van Baalen, C., Hoare, D. S., and Brandt, dim light. J. Bacteriol. 105:685-689. Venkataraman, G. S., and Neelakantan,

Heterotrophic Effect

growth of blue-green constituents growth

of the cellular on

nitrogen-fixing blue-green alga Cylindrospermum plants. J. Gen. Appl. Microbiol. 13:53-61.

muscicola

the root

Ward, W. R., Murray, B. C., and Malin, M. C. (1974). Climatic variations on Mars. 2. Evolution carbon dioxide atmosphere and polar caps. J. Geophysical Res. 79:3387-3395. Ward, W. R. (1973). Large scale variations in the obliquity of Mars. Science 181:260-262.

Weller, D., Doemel, W., and Brock, T. C. (1975). Requirement of low oxidation-reduction potential for photosynthesis in a blue-green alga (Phormid&m Sp.)Arch. Microbiol. 104:7 -13. Woiceshyn, P. M. (1974). Global seasonal atmospheric fluctuations on Mars. Icarus 22:325-344. Wolk, C. P. (1973). 37:32-101. Physiology and cytological chemistry of blue-green algae. Bact. Rev.

r_mA-L_ogloy,1_76

1 05

NATIC)NAL

AERCtNAUTIC5 WASHiNg,

AN[} TON. u D_

_f'A_

k 2054b

A[JMINibTRATI_)N

NATJON_.L

AERON,_UTtC_

AND

.................

..T, ,.,

,, .....

SPECIAL

---------FOURTH-CLASS BOOK

RATE

pos]Abl

45_ _ND

I[LS

PAIl,

POSTMASTI':R

If I'ndeliverahlt" l*l_st;ll 31alan41 )

"Map of the Planet Mars" by Percival Lowell, 1894

Courtes 7 Ioweli Obs_