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VOLUME LVII NUMBER 2

T H E

BOTAN ICAL GAZETTE

FEBR UAR Y i9"!

STUDIES ON THE REACTIONS OF PILOBOLUS TO

LIGHT STIMULI
HALiLY D. M. JOLIVETTE
(WITH TWELVE FIGURES)

The present investigationconcerns itself for the most part

with the problems of simultaneous stimulation. It was under-
taken with a view to settlingsome of the problemssuggestedin a
formerpaper on the light reactions of Pilobolus by Miss ALLEN
and myself(i). Some of the objectionsinherentin themethodsused
in theearlierworkwereeliminatedin the experimentsherereported,
and a studywas also made of the reactionof singlesporangiophores
toward the light.
The workwas begun underDr. R. A. HARPER at the University
of Wisconsin and completed under Dr. G. J. PEIRCE of Leland
StanfordJuniorUniversity. I wish to acknowledgemy indebted-
ness fortheircriticismsand suggestions. I wish also to expressmy
gratitudeto Dr. D. H. CAMPBELL forhis interestin my work and
forhis courtesyin extendingall the privilegesof his laboratory. I
thank Miss RUTH F. ALLEN, who began the study of the reactions
of a single sporangiophore of Pilobolus toward the light and
furnishedthe data taken on the evenings of May i8 and i9,
1910.
The present study of simultaneousstimulationby light is on
the question of directiveinfluence,and the literaturediscussedwill
concern that phase of light effect. The early work was carried
on with light and gravityas the stimuli. NOLL (4) in i8Q2 pub-
8g
go BOTANICAL GAZETTE [FEBRUARY

lished his theory of "heterogene induction." He reviewed the

earlierauthorson the subject and describedtheirconflicting results.
According to NOLL'S theory,if an organism is subjected to two
stimuli,one gives impetus to the other which carriesout the reac-
tion. One stimulusonly is responded to. There is no resultant
reaction toward the two stimuli. But when light and gravity
work together,thereis a change of geotonusdue to light.
Recently GUTTENBERG (2) studied the simultaneouseffectof
lightand gravity,using seedlingsof Avena sativa,Brassica Napus,
AgrostemmaGithago,and Helianthus. He used what he called the
compensationmethod. He tried different lightintensities. With
the higherintensitiesthe reactionwas toward the lightalone. By
gradually decreasing the intensities,GUTTENBERG found a certain
light strengthwhich just equaled that of gravity,and he obtained
a resultantreaction between the two. A little weaker or a little
strongerlight gave a resultantreaction, varyingaccordingto the
intensityof the light. GUTTENBERG considers this as evidence
against NOLL'S theoryof "heterogene induction."
RICHTER (5), workingalong the same lines as GUTTENBERG,
came to quite different conclusions. For his experimentsRICHTER
used Avena sativa, Vicia sativa, Vicia villosa,Brassica Napus, and
Helianthus. He followedGUTTENBERG'S method and in each case
carried on a set of experimentsin pure air and a similar one in
impure air. He concludes that GUTTENBERG did not establish a
resultantreactionbetween the effectof lightand gravityby means
of his compensationmethod,but that the latter's resultswere in-
fluencedby theimpureair in whichtheexperimentswereperformed.
GUTTENBERG (3) followedthis by a second paper in which he still
maintainedhis formerviews. In this he repeated his own experi-
ments,takingprecautionsto workunderpure air conditions.
The experimentson simultaneousstimulationreportedin this
paper were performedwith stimuliof the same kind, that is, they
were light stimulionly. Before enteringinto a descriptionof the
work of simultaneousstimulationof Pilobolus,an account of some
observationsmade on the reaction of a single sporangiophoreof
Pilobolus will be given.
I9I4] JOLIVETTE-PILOBOLUS 9I

Studyofthereactionsoftheindividualsporangiophore
to a singlelight
In the earlierexperiments(2), and also in thoseon simultaneous
light stimulation in this paper, I was concerned with a large
numberof sporangiophoresand with the net resultof the reaction.
This set of experimentswas inauguratedforthepurposeoffollowing
in detail the stages in the reactionof the individualsporangiophore
toward light. The horizontal microscopewas employed for this
purpose.
A cultureof Pilobolusin a 5-cm.flowerpot was used. The pot
was supported in an upright position. A thin glass Petri dish,
measuring5 cm. in diameterand 4 cm. in height,was placed over
the top of the flowerpot to keep the culture from drying. A
i6-c.-p. carbon filamentincandescent light was placed at a dis-
tance of 30 cm. fromthe culture,with the centralpoint of the fila-
ments 5 cm. above the level of the surface of the culture. The
experimentswere performedin the dark room and no other light
had access to the culture. A horizontal microscopewas placed
with the tube on a level with the surfaceof the cultureand at right
anglesto the directionof the lightrays reachingthe culture,so that
any bending toward the light could be observed. A micrometer
scale was placed in the ocular of the microscopein orderto measure
the change of position of the sporangiophore. In favorablecases
several sporangiophorescould be observed in the field of the
microscope.
The firstculture used in these experimentswas put in place
at 7:I5 P.M. The sporangiophorehad been exposed to the after-
noon lightand had grownstraightout towardit, makingan angle
of450 withthevertical. The culturewas placed withthesporangio-
phoresleaning away fromthe light,so that the angle between the
light rays and the sporangiophorewas about I350. At the time
whenthe experimentwas set up, the youngsporangiophores showed
no signsof the sporangialswellingor vesicularbulb.
Two sporangiophoreswere observed duringa period of 3 hrs.
on the eveningof May i8, i9io, and sketcheswere made at inter-
vals duringthe reaction. The exact timewhen the reactionof the
92 BOTANICAL GAZETTE [FEBRUARY

firstsporangiophorebecame perceptiblewas not determined. The

reactionwas distinctlyvisible at 7: 45, 30 min. afterit was exposed
to the light. So far as could be seen, the curvaturebegan at the
tip of the sporangiophore,the tip bending as it grew. The tip
distinctlystartedto curveupward at 7:45. This curve was some-
whatmorepronouncedat 7: 50. The tipwas vertical,havingmoved
throughan angle of 450. The radius of curvature was short. At
8:40 the tip had grownso that it was no longervertical,but made a
smaller angle with the incident light rays. At 9: 30 the tip had
grownaround so that it pointed in the directionof the light. It
had curvedabout I350 since the beginningof the observation. The
curvaturetook place as the growthoccurred; the curved end of the
sporangiophoreat this period was a well rounded hook. From

FIG. I

this time on the tip grew straight toward the light. The last
observationon thissporangiophorewas made at IO P.M. and showed
a pronouncedgrowthin the directionof the light.
Fig. i shows the stages that were sketched. The arrow indi-
cates the directionof the light. The bendingin thiscase had taken
place always at the tip, the growingpoint of the sporangiophore.
The older basal portion of the sporangiophoreappeared to main-
tain the formand position which it had at the beginningof the
experiment. If therewas any change,it was so slightas not to be
detected with the microscope.
The behavior of another sporangiophoreunder observation at
the same time was as follows. The reactionwas somewhatlonger,
no sign of curvaturebeing noticeableuntil 8:oo P.M., 45 min. after
the beginningof the experiment. The bendingprogressedslowly.
At 8:40 the tip had curved through450, the curve being gradual.
i0i41 JOLIVETTE-PILOBOLUS O0;

At 9: I 5 the tip seemed to have stopped bending and to have

grown straight. The angle made with the light was 60?. From
9: I5 to 9: 50 the increasein lengthwas slight,and the olderportion
of the curve seemed to have become slightlymorebent. At IO: I5
the terminalsporangial swellingwas well definedand the limits
of the vesicularbulb could be discerned. The directionof the tip
remainedunaltered and formedan angle of 6o0 with the incident
light rays (fig. 2).
A group of
sporangiophores
on the same cul-
ture as the two de-
scribed above and
subjected to the FIG. 2
same stimulation
from7: I5 P.M. until 9:55 were observedat 9:55. Three of them
were still turnedin the directionfromwhichthe afternoonlighthad
come and away fromthe lightused in the experiment. Apparently
there was no response toward the light stimulus. In these three
cases sporangium-formation had begun. The fourth sporangio-
phore had been subjected to the same conditions. The tip was
curved throughI350 and proceeded to growin the directionof the
light. The tip was still
slender and pointed.
The difference in length
between this one and
the other three was
noteworthy. It ex-
FIG. 3 needed the other three
by the length of the
portionbeyond the bend where it turned toward the light. The
sketches of the four sporangiophoresas they were at 9:55 are
shown in fig. 3.
At 8: I5, May i9, i9io, an olderculturewasused. Sporangium-
formationhad startedwhen the observationswere begun. At the
beginningof the experimentthe five that were chosen for study
stages, the youngestshowingthe sporangiumas a
were at different
94 BOTANICAL GAZETTE [FEBRUARY

small yellowknob, and the oldest having its sporangiumfullgrown

and turningblack. They were all pointed in the directionof the
afternoonlight,makingan angle of I35' with the incidentelectrical
lightrays. The fivewere watched closelyfrom8: I5 to 9:55 P.M.,
the position of the tip of each being observed and recordedevery
5 min. During this time their developmentcontinued normally,
the sporangia of the youngerones swelled, turnedgray, and then
nearly black, and the vesicular bulbs of all increased in size. In
no one of the fivewas thereany sign of bending toward the light,
althoughtheywerewatched fori hr. and 40 min.
On the eveningof May 2I, i910, observationswere made on the
reactions of three sporangiophores. The light was turned on at
7:38. The sporangiophoreswere inclinedaway fromthe direction
of the light at an angle of I25'. The -sporangialswellingin all of
themwas yellow. The vesicularbulbs had not begun to form. At
8:02 the sporangiophoreshad grown i mm. in length,but there
was no change of position due to the presence of the light. The
observationwas continued until 9:20 P.M. The sporangiophores
had not reacted toward the light, although they had continued
their normal development.
At 9:20 P.M., January i9, i9ii, observations were begun on
six sporangiophores. They were in the same field. The lightwas
turnedon at 7:20 P.M. weremadefor2 hrs. One
and observations
of the sporangiophoresstood vertically. It was slendertipped and
showed no signs of sporangial swelling. At 7:50, after 30 min.
exposureto the light,the tips showeda veryslightcurvature. The
reaction then stopped and the tip began to swell slightly. When
the lightwas turnedoff,the sporangial swellingwas distinct.
The remainingfive sporangiophoresat 7: 20 stood at an angle
of I350 to the incidentlightrays. All showed sporangialswelling,
but the vesicular bulb had not started to form. There were no
indicationsof response toward the lightin any of them,although
the sporangiophorescontinuedtheirnormal developmentthrough-
out the experiment. At the close of the experimentthe swelling
of the vesicularbulb on all of themwas just visible. Fig. 4 shows
a sketch of these sporangia as they appeared at the beginningof
the experiment.
I9I4] JOLIVETTE-PILOBOLUS 95

At 7:27, January 20, i9ii, the light was turned on a culture

which had been exposed to the afternoonlight,and observations
were begun on fivesporangiophoreswhichwere visible in one field.
The firstsporangiophoremade an angle of about 8o0with the direc-
tion of the light. It was slender
tipped. At 7: 58 the tip showed a
slight increase in length and this
portion was very slightly curved /
toward the light. At 8:23 there
was a slightincrease in curvature
towardthelight. At 8:42 thetip FIG. 4
had curved through350, making
an angle of450 withthe light. At 8: 50 the curvaturehad increased
so that the angle between the directionof light and the tip was
only 250. At 9:30 the tip was pointed directlytoward the light.
From this time the tip grewdirectlytoward the light (fig.5).
The second sporangio-
phore observed was in
much the same condition
at the beginning of the
experimentas the one just
FIG. 5 described, except that it
was about i mm. longer.
The reaction in this case was firstnoticeable at 8:oo P.M., the
curve being barely visible. The curvature then proceeded more
rapidly,and at 8:35 appeared more stronglythan in the othersin
the field. At this timeit had passed throughan angle of 30o. At
8:42 the angle traversed
was 450. At 8: 5o the
angle made with the
direction of the light
( M
rays was barely more FIG. 6
than ion, and at 9: I3
the tip was directedstraighttoward the light and it continuedin
that directionuntil 9:50, when the observationswere concluded.
The portion beyond the curve was then about three-fourths of
the lengthbelow (fig.6).
 96 BOTANICAL GAZETTE [FEBRUARY

The thirdsporangiophorewas inclined at an angle of 8o0 from

the incident light rays and measured 4 mm. in length. It was
slender tipped. Curvature was firstvisible at 7:58 and at 8:42
was still very slight. At 8: 50 therehad been a slight increase in
length,althoughno furtherchangein directionwas noted. At 9:40
the curvaturebecame more pronounced and at 9:50 the tip was
pointingalmost directlytoward the light. At this
time the curvature seemed to be arrested. No
\l\ \\l\ further
observations
weremade on thissporangio-
FIG. 7 phore(fig.7).
The fourth sporangiophore observed at the
same time stood at an angle of 8o0 with the directionof the light.
It was 2 mm. in lengthand the tip was slenderand tapering. The
reactionin this case was strikinglylike that of the thirdsporangio-
phore just described. They were very near together(fig.8).
The fifthsporangiophoremeasured i mm. in length at the
beginningof the experiment. It was slendertipped
and stood verticallyfromthe surfaceof the culture.
At 8:04 the tip had begun to curve. At 8:42 the
tip had curvedthroughan angleof 400. At 9:13 FIG. 8
it pointed in the direction of the light (fig. 9).
Observations were begun on two sporangiophores on one
cultureat 7:40, January2I, i9ii. Theywereslendertippedand
made an angle of about 50? withthe directionof the light. At 8: i8
both showed new growthwhich was curved slightly toward the
light. The curvature continued with the growth until at 8:50
the tip was directed straighttoward
the light. From that time until
9:50 when the observations were
FIG. 9 concluded, the sporangiophoresgrew
in the directionof the light (fig.io).
A group of fivesporangiophoreswas located in the fieldof the
horizontal microscope at 8:40 P.M., November2, i91 I . The spo-
rangiophoresshowed only slight differencesin length and were
inclinedat an angle of 250 fromthe vertical. They were placed so
that they leaned away from the light, making an angle of 95?-
The sporangialswellingwas just visible on the tips of all the spo-
I9I4] JOLIVETTE-PILOBOLUS 97

rangiophores. They were observed continuously,but therewas no

sign of a bending toward the light until 3:00 A.M. Meanwhile,
the sporangialswellinghad increasedin size and the vesicularbulb
was barely visible. At this time the sporangiophoreswere curved
slightlyat some distance below the sporangiumnearer the direc-
tion of the lamp. Observationswere not made again until 4:30,
when the sporangia were aimed directlytoward the light and the
vesicularbulb was distinct; the curvaturewas in the regionimme-
diately beneath the
vesicular bulb. At
6:30, whentheob-/I 'I1 f 1 {?
servationswere con- FIG. IO
eluded, the bulbs
had swollenconsiderably. The sketches(fig.I I) show the develop-
mentof one of the sporangiophoresof the group. The development
of all the others were remarkablyparallel with the one described.
Observationswere begun using a second horizontalmicroscope
on a sporangiophore at 9:oo P.M., November2, i9ii. The tip of
the sporangiophorewas just beginningto swell. The sporangium
was standingverticalto the surfaceof the substratum. The light
was placed at the angle above mentioned. The developmentof the
sporangialswellingcontinued,but no reactiontowardthe lightwas
visibleat II:35. Observations
weremadecon-
tinually until 3:00 A.M., when the sporangium
was rather well formedand the vesicular bulb
barely visible.
On the eveningof November3, I91I, two
FIG. II sporangia were observed for the firsttime at
IO:40. The first sporangiophorewas yellow
and rather blunt tipped, but as yet it did not show sporangial
swelling. It stood at an angle of about 200 fromthe vertical,thus
being inclinedaway fromthe lightat an angle of goo. There was
no signof a responsetowardthe lightat I2:00 P.M., but the tip
thenshowedthesporangialswelling. At 2:05 A.M. thesporangio-
phore began to curve towardthe light,the regionof curvaturebeing
then located at some distance below the sporangial swelling. At
2: I5 the curvaturewas more pronouncedbut still confinedto the
98 BOTANICAL GAZETTE [FEBRUARY

same region. The bendingcontinueduntil 5:30 A.M., when the tip

of the young sporangiumpointed directlytoward the light. The
entireportionbetweenthe regionofbendingand theyoungsporan-
gium was swollen slightly,showingthe beginningof the formation
of the vesicular bulb. At 6:30 the vesicular bulb was large and
turgid(fig.I2).
At the beginningof the observationthe second sporangiophore
in the same fieldof the microscopewas somewhatlongerthan the
firstand the sporangialswellingwas well formed. This sporangio-
phore also formedan angle of go9 with the light. The vesicular
bulb was not yet visible. At 2: I5 the sporangiophoreshad curved
throughan angle of 20? nearerto the directionof the incidentlight
rays. The curvaturein this case was also at some distance below
the sporangium swelling, and the space between the two was

FIG. I2

beginningto show signsof the formationof the vesicularbulb. At

5:30 A.M. the tip of the sporangiumwas aimed toward the light.
The vesicularbulb was still very inconspicuous. The bend in the
sporangiophorewas well roundedand had grownin lengthsince the
beginningof the curving. At 6:30 the vesicularbulb had swollen
so that it exceeded the diameter of the sporangiumby twice the
diameterof the latter. At 8:oo the developmentappeared fairly
complete. The sporangium was discharged between 9:50 and
Io:00 A.M. (fig. I2).
From the foregoingexperimentsthe followingconclusionsare
evident:
i. Growthtakes place at the tip in the young sporangiophore.
2. It is in thegrowingtip thatheliotropiccurvaturesare formed.

3. In no case has a heliotropiccurvaturebeen observed during

stagesin earlysporangium-formation.
I9I4] JOLIVETTE-PILOBOLUS 99

4. Sporangium-formation may start duringa reaction; in such

cases the reactionis delayed forsome time.
5. In case thereactiontowardlightisinterrupted bysporangium-
formation,it is resumedagain a littlebeforeor about the timewhen
the vesicularbulb is beginningto form.
6. After the sporangiumis formed,the growingand curving
portionis located immediatelybeneath the vesicularbulb.

The reactionof Pilobolus when exposed simultaneouslyto two

equal sources of light
The effectof exposinga cultureof Pilobolus simultaneouslyto
two equal sources of white light was studied. For this purpose
a light-proof box, measuringI 20 cm. long by 6o cm. wide by 6o cm.
high,was used. The box was made of pine and was painted a dull
black on the inside. Runninghorizontallyacross one end, 20 cm.
fromthe bottom,was an opening io cm. wide. Into the opening,
which was prepared with rabbeted edges, was introduceda gal-
vanized iron strip containingtwo openings i cm. in diameterand
9 cm. apart. The culture was placed at a distance of 25 cm.
from the central point between the openings and on a level
with them. It was placed with its surface vertical and facing
the side of thebox containingthe openings.
The object of the experimentdemanded that the lightfromthe
two openingsbe equal, but I know no methodsof obtainingabso-
lutely equal light intensities. We can only know that they are
approximatelyequal. In order to obtain as nearly as possible
equal illuminationat the two openings, one light of measured
intensitywas placed equidistant from the two openings of the
box and 40 cm. in frontof it. Two mirrorswere so adjusted that
the cultureinterceptedthe single spot of lightformedby the con-
vergenceof the two sets of lightrays. The lightfromeithermirror
was excludedfromthe opposite openingby means of the following
device. At rightangles to the edge of the box, along a line equi-
distantfromthe two openings,was placed an uprightpiece ofboard
measuring6o cm. high by 30 cm. wide by i cm. thick. At right
angles to thefirstpiece and parallel to the end of the box was nailed
a secondpiece measuring6o cm. highby io cm. wide by 5 mm. thick.
I0O BOTANICAL GAZETTE [FEBRUARY

Both of the pieces were painted a dull black. The surfaceof the
culturewas coveredwithblack paper, exposinga circulararea 2 cm.
in diameter. This small area was selected in order to exclude
objectionable features such as unevenness in surface of culture,
irregularityof distribution,etc. The number of sporangia was
thus somewhat limited,but the undesirable featuresabove men-
tioned were minimized.
This set of experimentswas carried on in a dark room at the
Universityof Wisconsin duringApril,May, and June i910, under
the directionof Dr. R. A. HARPER.
As previouslydescribed,a new set of sporangia matures daily
and is dischargedin the forenoonor early afternoon. The records
of the resultsof the experimentswere made daily in the late after-
noon or evening. A glass plate fittinginside the box and against
the openingscaught the sporangiaas theywere dischargedtoward
one or the other of the two lights.
The data were then recordedby means of a chart devised to
meet the requirementsof the experiment. The chartconsistedof a
large white sheet of paper divided by means of parallel lines into
vertical strips I cm. wide. This is the principleof the Wolfhiigel
counterused by bacteriologists,and it was well adapted to thework
in hand. The pieces of glass coveringeach of the i-cm. openings
fittedinto the i-cm. strips. In recordingthe data, the sporangia
fallingabove and below the openingin the i-cm. stripare recorded
with those strikingthe opening. This is entirelyfair,since,owing
to the object of the experiments,we are concernedonlywithlateral
distribution. Furthermore,our earlierexperimentsshowed clearly
the conditionsof vertical distribution. The data for these experi-
ments are recordedin table I.
In the firstexperiment86 sporangia were discharged on the
glass, 29 strikingthe vertical area containingthe opening to the
leftand 25 that to the right. In the second experimentthe total
numberwas 6o; 5 of thesewere on the area of the leftopeningand
20 on the right. In the thirdexperiment59 sporangiawerecounted
on the glass, 5 and i8 being found on the left and rightopenings
respectively. In the seventh all of the 22 sporangia discharged
werefiredtowardthe rightopening, io of themstrikingthe vertical
I9I4] JOLIVETTE-PILOBOLUS

area containing the opening. Experiment 26 shows a total of

142 sporangia,54 strikingthe vertical stripof the leftopening,30
that of the rightopening. Experiment27 shows 39 and 58 out of
a total of 204 sporangiaon the leftand rightopeningsrespectively;
and in experiment28, 334 sporangiawere discharged,84 on the left
and 78 on the rightopening.
Throughout the series of experimentssome of the sporangia
failed to hit eitheropening. In experimentI, 32 of the 86 spo-
rangia dischargedwere of thissort. The distributionon eitherside
of the two openings showed considerable variation. There were
9 sporangiain the i-cm. stripto the leftof the leftopeningand one
sporangiumin each of the next two strips. In the firsti-cm. strip
to the rightof the openingwere 6 sporangia. On the left side of
the rightopening therewere ii sporangia in the firststrip and I
in the third; 3 sporangia were foundin the firststripto the right
of this opening.
In the second (table I), 3 sporangiastruckthe glass withini cm.
to the leftof the leftopening. In the first,second,and thirdstrips
to the rightof the openingwere respectively4, 2, and i sporangia.
To the leftof the rightopeningwere 5 sporangia,all withinthe first
strip. The number of sporangia in the first three consecutive
stripsto the rightof the openingwere respectivelyi8, i, i. The
distributionand number of scattered sporangia in the remainder
of the experimentsshowed about the same degree of variation as
indicatedby the completedata (table I).
In the foregoingexperimentsthe number of sporangia which
strikethe openingsdoes not appear especially large. These alone
do not by any means give a complete conceptionof the accuracy
of the total dischargeof sporangia. A furtherknowledgeof the
distributionof these sporangia serves to correct the erroneous
impressiongiven by statingonly the numbersthat reach or do not
reach the opening. The accuracy is very striking,forfurthercon-
siderationshows that 29 of the32 sporangiathatmissedtheopening
in the firstexperimentwere foundwithinthe i-cm. stripson either
side of the openings; only 3 . 5 per cent felloutside that area.
A perusal of the remainingdata shows practically the same
accuracy of discharge. The greaterportionof the sporangia that
I02 BOTANICAL GAZETTE [FEBRUARY

failed to strikethe openingswerefoundin the stripsadjacent to the

openingsand only a very small percentagefelloutside this area.
It is plain fromthese data that although thereis much varia-
tion in the numberof sporangiafiredtoward the two openings,the
sporangiaclusterabout each of the openingsand show no tendency
to strikebetween the two. The sporangia are discharged,there-
TABLE I

86 .......i.. II 9 29 6!......... I II25 3.........I

6o...............3 5 4 21I........ 520 i8 I .... 2

56.. .... . .... .. I2 I.......2 2 514 141I32....... 4

25...............3 8 I12.. I.. I... I17......I...i 5
41...............2 9 4 2.....2 I I 410 4... II . 6
22...........................I ... 2 410 4... I.....7
35...............4 7 I1II1.1-I... 4 57 3..1I.... 8
45.........I I ..513 3 3.....I ... I III 5 .. .... 9
36............... 5 7 2.......I I 2 I 12 3 ..I I ... IO
54...............6 3 3..I .i....21I621 .....II

209.........I ..I 5 3 I 2 ..... . .....313.......... . 1I2

24 ............... 24 21I. .. I I .......II 2..1.... . 3
24 ...1............I9 3 ............. I8.. I.....i..14
14 ... ............I 4 ............. I 3 4 I.........I5
24............ . 1I3 7 2......... II 3 6 ..1I. ... i6
20...............2 3 3 ......... I 2 2 3 ..3.1... . 7
12................ . 4.. . ..2.. . .. I.. I 2 2. ........i8
20 ..1............I6 3 . I I.....I 5 1 1... 19.....I
7 . ................2 2 ... ......I.....2 ... ........20
35 .1...........I310 5 2.........34 7.......... 21
27...............2 8 2 4 ......... 2 3 5 I.........22
14.................5 I I... ........... 4 3.........23
22 ..1............I4 3.. ...2.......... 7 5..........24
34...............612 5 ....... I.. ...26 I I.......25

I42.........I 4 16 54 17 I I ... I .... 0

O30 6 I.......26
204.........2 3 4 23 39 96 3 42I 2720 58204.......27

334 ..1..1.II2 32484 23 54 26 3 934 7839 82 3 328

I33.......I.....2 8431I22.....I 3 61i632 4 2.1..I..29
178.......I 2 I 8 26 52 28 6 I I 2 I 2 16 21 8 I.....I 30
I46......... I.. 41I225 8 42 2.. 3 427 44 7..I..1.1I31
24 ............. 1 2 2 I ... I I I.....4 9 I I.......32
252 ..1 2 3 23 5 2525 i6 8 5 45 512 22 4433 9 986 33
6i.. .... . ..2 2 52 53 4 21I 147113 4 32 I1...34
30.......I 2 I I132 2.....I 3 42 3 2 211 I .......35
I76 .1.....I251I71I59 6.. 3 ... 81I325 20 19 13 5 8 4 3 36

fore,towardone or the otherof the two lights. In otherwords,as

shown by the earlier set of experiments,there is no resultant
reactiondue to the presenceof the two lights. If therewere, the
sporangia would be for the most part between the two openings.
that this simple organism,when subjected
It is clear, therefore,
simultaneously to two equal light stimuli,will respond to one of
the two stimuli,to the completeexclusionof the other.
I9I4] JOLIVETTE-PILOBOLUS I03

The sporangia, however,are not dischargedin equal numbers

towardthe two openings. In the extremecase all mightgo to one
or the other of the openings. It happened that in only one case
(experiment7, table I) did this occur, and then the total number
was 22, the number being small and affordingless chance for
variation than would a larger number. The entire data thus
show a great diversityof resultsas regardsthe numberfiredtoward
eitheropening. It is significantthat the elementof chance enters
stronglyinto these results.
The fact that the sporangia do fire toward one or the other
opening and in any degree of variation suggests further that
althougha large numberof sporangiophoresarise fromone myce-
lium, they are not gregarious as regards physiological response
toward light stimuli, that is, they act as separate individuals
toward a light stimulus.
A furtherset of experimentswas arrangedto determinewhether
the sporangia to the rightof the cultureare dischargedtoward the
rightopeningand those to the leftof the cultureto the leftopening.
With the apparatus arrangedas before,a second seriesof experi-
ments was made with a thin plate of glass placed with its edge
vertical to the middle line of the exposed surfaceof the culture.
This glass would then interceptthe sporangia dischargedfromthe
leftside of the cultureto the rightopeningand vice versa.
In the firstexperiment2 sporangia struck each of the two
openings,the numberof sporangia in the consecutive i-cm. areas
to the left of the left opening are 8, 6, and I; 3 sporangia were
found on the i-cm. area to the rightof the opening. The first,
second, and third i-cm. areas to the leftof the rightopening con-
tained respectivelyI, 2, and i sporangia. The glass placed verti-
callyat rightangles to the surfaceof theculturereceived7 sporangia
on the leftside and I 5 on the right. The distributionof those on
the leftwas as follows: 4 in the ist cm. towardthe culture,2 in the
4th cm., and i in the 7th. The distributionof the sporangia on
the right-handside of the glass shows 7, 5, and 3 in the first3 cm.
These data, togetherwith those of the second and thirdexperi-
ments made in this series, are found in table II. These experi-
mentstend to show that some of the sporangiaon the rightside of
 I04 BOTANICAL GAZETTE [FEBRUARY

the culturefiretoward the left opening and vice versa. Some of

thismay be due to reflectionfromthe glass.

TABLE II
The culture midway between and facing the two openings; a glass plate before the openings; a second
glass plate at right angles to the surface of the culture. The data fromthe plates before the openings are given
in the firstpart of the table; those fromthe plate at rightangles to the surface of the culture are referredto by the
marks *, t, and t.

i. Date

26 _ ..- .. .. I 6 8 2 3 ... ... ... ... I 2 I 2 ... ... ...... ... ... I* 6/I5/'IO
7:30 P.M.
7 ... ... ... ... ... ... I ... ... ... ... ... 2 ... ... I 2 I ... . . ... ... . . . 2t 6/i6/'io
7:35 P.M.
38 ... ... ... ... ... ... I 4 7 9 ... I . .. 2 7 I 4 I . .. ... 3 6/,7/'IO
7: 20 P.M.

* Sporangiadischarged on glass at J4.. ... 2 ... ... I ... ... Surfaceofglass to leftofculture
right angles to surface of culture 7 5 3 ... ... ... ... ... ...
it right "

f Sporangia discharged on glass at f6 3 ... . .. ... ... ... ... . ... left "
right ""
I it it it
right angles to surface of culture ',2 I 2 I I ..". ... ...

t Sporangia discharged on glass at f " " "it "

left "
right angles to surface of culture l I . . . 2 I .. . ..... it it
right ""

The behavior of Pilobolus when given two equal sources of light

but withthe angle between the two sets of incident
lightrays varied
The box used for this set of experimentswas made of redwood
and measured I20 cm. long by 45 cm. wide and 35 cm. high. A
hingedcover was so rabbetedas to be light-tightand was supplied
with ice box catches in order to fasten it down tightly. At one
end of the box, I4 cm. fromthe base, was an opening 9 cm. wide
all the way across fromside to side and so arrangedwith galvanized
iron rabbets as to carry strips of galvanized iron io cm. wide
whichjust closed theopening. These stripscontainedthe openings
throughwhich the light was admitted to the culture. They were
made of galvanized iron so that they would be thin enough to
avoid shadowsbeing cast by the rimof theopenings. The openings
were I cm. in diameter and their centerswere the followingdis-
tances apart: I cm., 2 cm., 3 cm., etc. The inner surfaceof the
slide containingthe openingwas flushwith that of the end of the
I9I4] JOLIVETTE-PILOBOLUS I05

box, so that the glass plate on which the spores were caught fitted
closely against the slide containingthe openings. The openings
not desiredcould be closed by means of two additionalslidesfitted
in the groove fromeitherend and outside the slide containingthe
openings. The inside of the box and of the stripswas dull black.
The culture in this series was kept throughoutthe experiment
at a distance of 23 cm. from the central point between the two
openings. The surfaceof the culture was vertical and faced the
slide containing the two openings. The exposed surface of the
culture was 2 cm. in diameter. The light enteringeach of the
openingswas made to fall upon the exposed surfaceof the culture.
The intensitiesof the light enteringthe two openings were equal
or as nearlyso as theycould be made by measurement. The follow-
ing device was followedin orderto make the lightsas nearlyequal
as possible. A singlecarbonfilamentincandescentlamp was placed
at a distance midway between the two openings. By means of
two mirrorsplaced at equal distancesand equal anglesone on either
side of the lamp, the lightwas reflectedthroughthe openingto the
exposed surfaceof the culture. In order to exclude the light of
eithermirrorfromthe opposite opening a partitionwas set up in
frontof the box midwaybetweenthe two openings,exactlysimilar
to that used in the set of experimentslast described.
With the change in the distance between the two openings it
was necessaryto change slightlythe angles of the mirrorsfromthe
light in order that the spot of light reflectedfrom the mirrors
throughtheopeningswould striketheexposedsurfaceofthe culture.
The conditionoftheexperimentsthusnecessitateda slightchangeof
lightintensityfromexperimentto experiment,but the intensities
of the lightpassing throughthe two openingsat any one time were
equal.
This set of experimentswas performedin a dark room in the
botanical laboratoryof Leland StanfordJuniorUniversity.
In the firstexperiment,when the distance between the centers
of the two openings was 5 cm., a total number of 38 sporangia
werefiredon the glass, io and I3 strikingthe leftand rightopenings
respectively. Only 2 of the I5 sporangiawhichfailed to strikethe
openingwere outside of the adjacent i-cm. strips.
io6 BOTANICAL GAZETTE [FEBRUARY

In the second experiment,with the centerof the openings4 cm.

apart, i9 sporangia struckthe glass; io were on the leftopening,
7 on the right. Of the remaining2 sporangia, i struckin each of
the i-cm. stripsto the leftand rightof the leftopening.
The thirdexperimentshows a total dischargeof 63 sporangia.
Of that numberi6 struckthe leftopeningand 20 the rightopening;
27 failed to reach eitheropening,but all of that number,excepting
3, were withini cm. of the opening.
When the distance between the openings was I4 cm., 50 spo-
rangia weredischarged,i5 strikingeach oftheopenings; 6 sporangia
were counted within the firsti-cm. strip to the left of the left
opening,and i in the thirdstrip; 3 and 2 were withinthe firstand
second stripto the rightof that opening; 5 sporangiastruckwithin
the i-cm. strip to the leftof the rightopening,and 2 and i in the
firstand second stripson the right-handside.
With a distance of 27 cm. between the two openings, 200 SpO-
rangia weredischargedon theglass,6i and 37 strikingtherespective
openingsto the leftand right. The majorityof scatteredsporangia
are again grouped around the two openings. The data for these
experimentsare tabulated in completeformin table III.
The resultsof the experimentsin which the distances between
the openingswerevaried agreewiththoseobtainedin the preceding
set of experiments,where the openingswere kept at the same dis-
tance throughoutthe set of experiments. The sporangia were
firedwith great accuracy towardone or the otherof the two open-
ings. The distributionof the sporangia about the openingsvaried
in about the same degreeas in the case just mentioned; and in the
same way, the sporangiawhich are outside the vertical stripscon-
tainingthe openingsare mostlywithini cm. of one of the openings.

The reactionof Pilobolus when stimulatedsimultaneouslyby

lightsof different
wave-length
The problemsconnected with the simultaneousstimulationof
an organismby lightrays of various wave-lengthsofferan interest-
ing fieldto the investigator. But I know of no way of accurately
comparing lights of differentcolors as to the total amounts of
19I4] JOLIVETTE-PILOBOLUS I07

~ inIt lo -O 0O% H N in l

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

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

. . . w. . . . .

ZN~~~~~~~
.. . . . . . . *.*.

... H . ...

. . . ',. . . . . . .

.
0 _ _ . _ _ _ _.__C,_0
_ _ . _ *_ *_ _ _ _
.* N

PA

Nd . H % . . . .

I0

0 . . ,

vZ . . . . . . . . . . . . o. c .

00C, ('n : :: :::0'tHHMc0

H ~~~~~~~~~~ 0 . . ...... uY

0
O)
H .....
.......
** 0vO N t
N0
H .
..
H H
..
*

~~~~~ NA 0N NQ0 m0 0 NH

0 t-~~~~~~~'t0Co-(', 0i
io8 BOTANICAL GAZETTE [FEBRUARY

radiant energy. The followingexperiments,therefore, are qualita-

tive only; but I hope theywill prove suggestive.
The differentcolorsof the spectrumare representedin different
proportionsin the various incandescentlamps. These filaments
are of standard make and the energyof the bulbs is measured in
candle-powers. Thus by using bulbs of equal candle-powerand
currentof known intensitywe shall have somewhat comparable
quantities.
Althoughthis method is not all that could be desired,it has a
very marked advantage over the colored solutionsused by SACHS,
and also the monochromaticglass plates that are so generallyused
in workon the effectsof raysofvarious wave-lengths. The colored
solutions absorb a large portion of the total energy emanating
fromthe source,different solutionsdifferingin thisrespect. In the
case of the solutions (and it is also true of the colored plates)
neitherthe intensitynor the energyof the lightscan be compared.
The incandescentlamps offerat least the advantage that they are
of nominal commercialvalue; and with the advent of the knowl-
edge of methodsof comparisonof intensitiesof coloredlights,some
parts of theirspectrum
exact idea of the intensitiesin the different
may be obtained. Some study has been made to determinethe
distributionof the different wave-lengthsin the different incandes-
forthe presentinstance. It
cent lamps, but so farit is insufficient
is known,however,that of the threeincandescentlamps used in the
experiments,the tungstenhas the largestproportionof the actinic
rays, the tantalum next, and the carbon least. The results with
thesefilamentsmay thus serve to checkup withthose of the earlier
work,in which the solutions and the plates of colored glass were
used.
Experimentswere made in orderto test the relativeefficiencies
of differentincandescentlamps in bringingabout the reactionof
Pilobolus. In these experimentsthe carbonfilament,the tantalum,
and the tungsten were compared. The experimentswere per-
formedin the dark room,usingthe redwoodlight-proof box already
described. The openingswere i cm. in diameterand the distance
between them fromcenterto centerwas io cm. The culturewas
placed 23 cm. from the point midway between the two openings.
I914] JOLIVETTE-PILOBOLUS I09

The entiresurfaceof the culture,whichwas 5 cm. in diameter,was

exposed. Before each of the openings, at equal distances, was
placed one of the two lightsto be tested. The angle at whichthe
lamps were placed was such that the spot of lightfromeach lamp
strucksymmetrically the open surfaceof the culture.
The series of experimentswas begun with a 32-candle-power
carbon filamentlamp beforeone opening and a 20-watt tungsten
beforethe other. The firstexperimentgave a total of 92 sporangia;
3I sporangia were dischargedtoward the carbon filamentand 6i
toward the tungsten. In the second experimenti63 sporangia
were discharged; 68 toward the carbon filamentand 95 towardthe
tungsten. In the thirdexperiment387 out of a total of 784 spo-
rangiawent to the tungsten. Of the ii i6 firedin thefourthexperi-
ment, II3 went to the carbon, I003 to the tungsten. The next
experimentshowsa total dischargeof 228 sporangia; 78 towardthe
carbon, i50 toward the tungsten. A new carbon lamp was put in
the place of the old in the seventh experiment,but again a very
much largerpercentagewas aimed toward the tungsten; 589 spo-
rangia were fired,the ratio standing i68 toward the carbon as
against 43I toward the tungsten. The data for this set of experi-
ments are given in complete formin table IV. From this table
a comparisonof the accuracy of aim of Pilobolus towardthe carbon
and tungstenlamps can be made.
TABLE IV

Total

92 ... ... .. ... . ... 6 I9 6 ... ... ... . . ... 3 7 42 9 ... . . . . ..

163 ..........I 3 4714 21I.... II 3 I5 54 20.... I....
784. I I . 6 64 203 86 I5 6 4 2 2 5 8440253 8 3 I. .
iii6 ... ... ... ... ... 3 i96919 3 ... 2 2 3 5 I7I70 607 I79i8.
228 ... .1.5. .4.. ... ... ... I7 I24 .4 6 22 IOI I7 ... . . . .
894 ... ... ... ... I ... 39 I I I 2I 7 I I ... 2 2 9 58 503 I36 4 I ... ... ...
589 .. ., _ I 7 48 78 24 3 4 2 3 6 8 8 69 258 63 4 I 2 .

Of the 3I sporangia discharged toward the carbon lamp in

the firstexperiment,i9 (6i per cent) struck the glass over the
i-cm. vertical strip containingthe opening. Of the 6i sporangia
iio BOTANICAL GAZETTE [FEBRUARY

discharged toward the tungsten lamp, 49 (8o per cent) struck

withinthe correspondingstrip.
In the second experiment,of the I63 sporangia discharged,
68 were dischargedtoward the carbon filament,47 (69 per cent)
strikingthe i-cm. stripovertheopening; 95 weredischargedtoward
the tungsten,54 (56 per cent) on the strip over the opening. In
this experiment,unlike the first,the largerpercentagestruckthe
stripover the openingin the case of the carbon filament.
In the thirdexperiment,387 sporangiawere dischargedtoward
the carbon, 203 (5I .7 per cent) of them in the vertical strip con-
taining the opening; 397 were discharged toward the tungsten,
253 (65.2 per cent) on the regionof the opening.
The fourthexperimentshows 69 sporangia,whichis 6i per cent
of the II3 sporangia discharged,toward the carbon, on the i-cm.
strip containingthe opening; and 607 (6i per cent) of the I003
sporangiafiredtowardthe tungstenon the same region.
The remainingexperimentsof this series all show greateraccu-
racyin the tungstenlightthan in the carbon filamentlight. With
one exception,that of the second experiment,the dischargeof the
sporangiais more accurate toward the 20-watt tungstenused than
toward the 32-candle-powercarbon filamentlamp, although the
energyof the tungstenlamp is only half that of the carbonlamp.
Again, the percentageswhich strikethe openingsprobably do
not at firstglance appear remarkable,but on noting in the first
experimentthat the I2 sporangiathat did not strikethe openingin
the case of thosefiredtowardthe carbon lightwere all withini cm.
of it, the accuracy is striking. Of the i9 sporangia which failed
to strike the opening before the tungstenlamp, i6 were within
i cm. of the opening and the remaining3 were within2 cm. of the
opening.
In the second experiment,2I of the sporangiafiredtoward the
carbon lightmissed the verticalstripcontainingthe opening; I7 of
that number were withini cm. distance of the strip. Of the 4I
sporangiathat missed the openingin the case of thosefiredtoward
the tungsten,35 struckwithin i cm. of the opening.
In the remainingexperiments,most of the sporangia which
failed to reach the opening in the stripsfell withinthe firsti-cm.
I9I4] JOLIVETTE-PILOBOLUS III

strip on either side of them (table IV), showing the remarkable

precisionwith which the sporangiumis throwntoward a light.
A seriesof experimentswas made to test the relativeefficiencies
of a tungstenand a tantalumlamp in bringingabout a reactionof
Pilobolus; 40-watt lamps of each kind were used. The tungsten
was placed beforethe left opening, the tantalum beforethe right
opening at a distance of 40 cm.
TABLE V

Total tE
H Hd

373 ... ..II .. I 37 56 25 I0 I 3 3 2 6 I3 3 I I7 5 I ... ... ...

628 ... ...... ... I4I5 I00243 I0032 5 454 510O 34 34 I73 3
206 ... ... ... . . . 24 7I I34 I 3 4 5 3 4 i839 I 2 ... ... ... ..
32I ... ... . . . 7 36 I04 42 5 ... I ... ... 2 3 4o 6i i62 ... ... ... ...
i85 ...........9 25 6123 6 5...I 2 I1415 24 7 2........
i66 . 2 I 8 25 5 52
2667 I ... ... ... I 9 I3 3 1. 2
688 ... ... ... ... 3 20 II2 29I I02 i9 6 2 2 2 2 5 29 47 24 ... ... ... ... ...
886 ... .. .5.0.. ... ... 176 363 I585I 20 II 4 2 3 I I422 9 I I ... ... ..
574 ... ... 3 5 I5 69 248 53 i82 I
.1..... I 5 24 43 23 2 I I ... ...
...
I09I ... ... ... 6 20 46 I56 354 i64 46 2i 878 I320 40I4I 5I I 55
464 ... I I 5 2 2i 8i i66 7I 29 I73 2 3 3 8 7 27 9 4 2 2 ...
367 .. ... ... I I782 143 28 25 I5 I 7 I_ 4 8 27 7 I
_.. ... . .

In the firstexperiment,373 sporangia were dischargedon the

glass, 294 toward the tungstenand 79 toward the tantalum. The
second experimentshowed a total numberof 628 sporangia, with
516 and II2 toward the tungstenand tantalum respectively. The
numberof sporangia dischargedin the thirdexperimentwas 206;
of these, ii8 were firedtoward the tungstenand 88 toward the
tantalum. The total number of sporangia in the fourthexperi-
ment was 32I, I97 and I24 being firedtoward the tungstenandS
tantalum. In the fifthexperiment, i85 were discharged, I30'
toward the tungsten and 55 toward the tantalum. The sixth
experimentalso showed a largernumberhad been firedtoward the
tungsten. The data for this set of experimentsare found in
table V.
The accuracy of aim toward the tungstenand the tantalum
lamps was compared as in the precedingexperiments. In the first
experiment 46.6 per cent weredischargedtowardthe tungsten,3 . 2
per cent toward the tantalum. In the second experimentthe
II2 BOTANICAL GAZETTE [FEBRUARY

percentagesfiredtoward the tungstenand tantalum were respec-

tively 47 and 33.5. The third experimentshows 6o. i per cent
beforethe tungstenand 44.3 beforethe tantalum; the fourth,52.7
and 49. i respectively;the fifth,46.9 and 43.6; and the sixth,48.9
and 44.8. The tenthand twelfthexperimentsshow slightlylarger
percentageson the openingbeforethe tantalum. The percentages
figuredout for the numbers strikingeither opening are given
in table VI. The largerpercentagesstrikethe openingbeforethe
tungstenin these experiments. Most of the sporangiamissingthe
opening strips in these experimentswere found in the adjacent
i-cm. strips,as was the case in the foregoingexperiments.
TABLE VI

Number Percentages Percentages Number Percentages Percentages

of before before of before before
experiment tungstenlamp tantalumlamp experiment tungsten
lamp tantalumlamp

I........... 46.6 39.2 7. 44.3 42.7

2......... 47 33.5 8........ 4I.3 40
3......... 6o.I 44.3 9...... 59.5 43
4......... 52.7 49I IO .42.9 47.6
5......... 46.9 43.6 II .4I.7 40.9
6........ . 48.9 44.8 I2 .44.2 56.7

In the next seriesof experimentsa 40-watt tantalumwas placed

beforeone openingand a 20-watt tungstenbeforethe other. The
total numberof sporangia firedin the firstexperimentwas Io04;
73I were on the half of the glass toward the tantalum and 273
are on the half toward the tungsten.
The second experimentshowsa total dischargeof407 sporangia,
329 on the tantalumas against 78 on the tungsten. The thirdand
fourthexperimentsboth show larger numbers on the tantalum.
The data forthis set of experimentsare foundin table VII. The
accuracyofaim towardthe two lightsmay also be obtainedfromthe
data in this table. Of the 73I sporangia dischargedtoward the
tantalumlamp in the firstexperiment,420 (54 per cent) struckthe
i-cm. strip containingthe opening. Of the .273 firedtoward the
tungsten,I49 (54 per cent) were on the correspondingstrip. In
the second experiment,the numberof sporangia firedtoward the
tantalumlamp was 329; of these i6i (49 per cent) werein line with
the opening. The total toward the tungstenwas 78, with 37
 I9I4] JOLIVETTE-PILOBOLUS

(47 per cent) on the opening strip. In the thirdexperiment,the

ratio of the percentagesof those on the i-cm. stripcontainingthe
opening,in thecase of the tantalumlamp, to thaton thecorrespond-
ing stripin the case of the tungstenis as 57 to 6i. In the fourth
experimentthe accuracy of aim is very nearly the same toward
the two lamps, although a greaterproportionof the total number
favorthe tantalumlamp. The comparisonof a 40-watt tantalum
lamp against a 20-watt tungstenshows that a larger number of
sporangia are discharged toward the tantalum, but that there is
very little differenceas regards the accuracy of aim toward the
two lamps.
As in the set of experimentslast described,a close examination
of the data (table VII) reveals the fact that most of the sporangia
that missed the verticalstripscontainingthe openingswere found
withini cm. of them. An average of 7.8 per cent struckthe glass
more than i cm. laterallyfromthe opening.
TABLE VII

Total c
0 0

I004 . ... ... ... 4 9 I37420 II0 3III 5 105 3 I240 I49 55 3 ... . . .
407 ... ... ... ... ... 3 89 i6I 49 I27 5 5 5 3 3 I537 I3... ... ... ... ...
I25 ... ... ... ... ... ... i8 40 IO . . . . . . . . . . . . . . . . . .II 34 10 ... ... ... ... ..
292 ... ... I... 4 35 III 34 I0 3 ... I I 4 5 23 45 15.. ... ... ... ...

A 40-watttantalumlamp was next comparedwith a 32-candle-

power carbon filamentwith the resultsshown in table VIII. The
firstexperimentgave a total of 96 sporangia,36 of whichwere dis-
charged toward the tantalum and 6o toward the carbon. The
second experimentgave a total of 23I sporangia,with i64 fired
toward the tantalum and 67 toward the carbon. In the third
experiment365 sporangia struckthe glass, 268 over the tantalum
and 96 over the carbon. In the fourthexperiment,of the I57 SpO-
rangia fired,98 were on the glass over the tantalumand 59 over
the carbon. The fifth,experimentshoweda total dischargeof I039,
674 being found on the glass beforethe tantalum,465 beforethe
carbon. In the remainderof the experiments,as can be seen from
 II4 BOTANICAL GAZETTE [FEBRUARY

the data (table VIII), there is a larger percentage of sporangia

over the tantalum. Thus, with the exceptionof the firstexperi-
ment,the numberdischargedin the directionof the tantalumlamp
surpassed that dischargedtoward the carbon.
TABLE VIII

Total

96 .............8 23 5......... I I0 40 9 ...........

23I1.............43 7934 6 I I.....3 21I7 321I3...........
365 .............65136 491I5 3 I.......1I28 481I4 5.........
I57 .............245714 2 I ......... I 6 40101I... I....
I039 ........... 7IO63231I07 15 3 2 2 2 I15 91 2421I05 8.........
76.........1I212 2511 3........ 2 61I76...........
404 ...........I 35 164 29 I.......2 I I 26 110 31 3.........
I775 I II 2 3361i64748I70 31o61I 511I2610o6283135 20105 I
100 .............7 47 III...........I 6 23 4...........
507.........1I5 561I54 59 521211 I5 51I2140 3.........
1442 .. ...I3 1 8208 55324624 4822 7 628 8518363 6 2 3 . 2
8I3 5/11 44 IO15 20 58I78 77I5 I5I7 i6I8 25 3i65 93 711818 9 511 /4
174......... 13 29 44 82111 I3 2 71I53022 7.........
249....... I .. 3 20 951811I.. .... 13 30581I71I.........
III .............II 69 17 I.............2 9 2...........
104 .10 . . ......I 6i 1I7 I.............3 I0 2...........
184.1910119... .... g OII I............. 7 28 9...........
I94 .............2262 27I.............6 5 520 I.........
I23 .1. . . . . . 2 35 8.............2 10 42 14.....II
I23I1....... 2 14 40118 2661i62 40 I8 2 2 I 2 501I22 2571I07 42 9 3 .. I
44.. . .. . .. 5 19 4 I ... I.......I I II I...........
43 ...........I 5 i6 .............I... 2 12 6...........
561 ........... 6 98 I46 35 5 5 5 2 I 2 8 56 I22 44 7.........

Of the 36 sporangiadischargedin the directionof the tantalum

lamp in the firstexperiment,23 (63 .8 per cent) struck the glass
withinthe i-cm. stripcontainingthe opening. Of the 6o sporangia
firedtowardthe carbon,40 (66.6 per cent) struckwithinthe i-cm.
strip containingthe opening. In the second experiment,79 (48. 2
per cent) of the 164 sporangia that are discharged toward the
tantalum are found in the i-cm. strip-overthe opening; 32 (48
per cent) of the 67 dischargedtoward the carbon are on the corre-
sponding strip. In the third experimentthe percentages which
strikethe glass on the i-cm. verticalstripover the openingbefore
the tantalumand carbon filamentlamps are respectively50.7 and
50. In the fourthexperiment58 per cent of the sporangia fired
toward the tantalum and 67.7 per cent of those firedtoward the
carbon filamentlamp strike the strip containing the respective
openings.
I9I4] JOLIVETTE-PILOBOLUS

The above percentagesstrikingthe area of the i-cm. opening

before each of the two lamps compared are given,togetherwith
those computedforthe remainingexperiments,in table IX. The
complete data fromwhich the percentageswere figuredare found
in table VIII.
TABLE IX
PERCENTAGES OF THE TOTAL NUMBER OF SPORANGIA DISCHARGED TOWARD THE
TANTALUM AND CARBON FILAMENT LAMPS WHICH STRUCK THE I-CM.
VERTICAL STRIPS COVERING THE OPENING IN EACH CASE IN
EXPERIMENTS I-23 IN TABLE VIII

Number Percentage Percentage Number Percentage Percentage

of before before of before before
experiment tantalumlamp carbonlamp experiment tantalumlamp carbonlamp

I.......... 63.8 66.6 I3 ......... 49.3 34.8

2.......... 48.2 48 I4 ......... 68.3 52.7
3.......... 50.7 50 I5 ......... 60.2 69.2
4........... 58 67.7 i6 ......... 68.5 66.7
5.......... 68.i 52 I7 ......... 78.5 63.I
6.......... 55.5 58 i8 ......... 55-3 67
7.......... 7I.2 63.2 I9......... 63.6 6i.8
8.......... 63.8 47 20 ......... 32.6 43.I
9.......... 7I.2 67.6 21 ......... 63-3 78.5
10.......... 57 54.2 22 ......... 72.7 52.3
II .......... 52.4 47.6 23 ......... 45.6 50.4
12........ . 4I.7 24.2
Average. . 58.6 55.9

In this set of experimentsthe accuracy of dischargevaries con-

siderably. In some cases the discharge toward the tantalum is
more accurate; in othersthat toward the carbon lamp is more so.
On the whole, however, the discharge is a little more accurate
toward the tantalum,but the difference is so small that it is prac-
tically negligible.
From the percentagesin table IX, it stands out clearly that
whereverthere was a small percentagewhich struck the opening
beforeone lamp, therewas usually a comparativelysmall percen-
tage on the correspondingarea before the second light. In the
firstexperiment,63.8 and 66.6 were the percentagesstrikingthe
two openings. In the second experiment,48.2 and 48 per cent
struckthe openings. In some cases thereis less uniformity. The
eighth experimentshows 63.8 per cent on one opening and 47
per cent on the other. But on the whole, a small percentageon
ii6 BOTANICAL GAZETTE [FEBRUARY

one openingis usually accompanied by a correspondingpercentage

on the other. It mightbe suggestedthat this may be due to the
generalconditionof the cultureat the timeof the experiment. The
accuracymightbe affectedby food supply,moisture,temperature,
and other factorsof importanceto the physiologicalconditionof
the sporangiophoresat the time of stimulation.
Here, as in the previous experiments,the sporangia that strike
outside of the openingare to be foundforthe most part in the first
verticalstripsto the leftand rightoftheopenings. An examination
of the data (table VIII) will show a good proportionof the cases
where all that were firedtoward a single light are found on the
opening or adjacent strips. Thus it is clear that the accuracy is
much greaterthan it would appear froman examinationof table IX
alone.

Summary
Physiologists,in studying the reactions of plants to stimuli,
have for the most part worked with phototactic organisms or
organismsof considerablecomplexity,individuals in which there
was a differentiation of tissues, where the cells in one portion of
the body may receive a stimulus, another perceive it, and still
another respond to it. Such a study has the disadvantage of
dealing with too many factors and accompanying phenomena.
In Pilobolus the reactionis marked and can be easily studied. A
singlecell receivesthe stimulusand respondsto it. The protoplasm
of the cell receives the stimulus,perceives it, and reacts. The
accuracy of response of Pilobolus toward the lightis remarkable,
whenwe considerits size and the distance throughwhich it throws
its sporangia. The sporangiophorescarcelyever exceeds i cm. in
length,and is usually somewhatshorter,whilethe distancethrough
whichit dischargesthesporangiumin mostoftheexperiments is over
25 times that measurement. The accuracy of response and the
nicetyof organizationof such a mechanismcan well be appreciated
fromthestudyofsuch experiments. From suchworkthe capacities
of a single cell can best be realized.
The resultsof the experimentsin whichPilobolus is stimulated
simultaneouslyby two lights bear directlyon NOLL'S (4) theory
I9141 JOLIVETTE-PILOBOLUS II7

of "heterogene induction." According to NOLL, the reaction of

an organismto one of two stimuliexcludes the effectof the second
stimulus. His workwas concernedwithtwo verydifferent kindsof
stimuli,light and gravity. The reaction to the stimulusof light
excludedany responseto the stimulusof gravity. Recent workers,
GUTTENBERG (2) and RICHTER (5), have interested themselves
along the same lines. GUTTENBERGmaintains that if the light
stimulusbe diminishedsufficiently, a resultantreaction between
light and gravitywill occur; that NOLL and the earlier workers
had used light that was too intense. To RICHTER'S (5) criticism
that his resultswere due to impureair, GUTTENBERG (3) responded
by furtherworkunderimprovedconditionsand reached essentially
the same resultsas before.
In experimentson simultaneous stimulation of Pilobolus by
lightsof the same kind or of differentkinds,we are dealing,unlike
either of the foregoingcases, with simultaneous stimuli of one
kind, namely, light alone. It was possible to have the stimuliat
least approximatelyequal, and it was possible to have the arrange-
ment such that neithersource of stimulationhad any advantage
over the other. Further,the organismworked with was a simple
one, the reactionconcerningonly a singlecell. And the net result
of the reactionwas shown so plainly in the distributionof the dis-
charged sporangia that it seems impossible that any indefinite-
ness or uncertaintycould be entertainedas to the reaction. The
sporangia clustered always about one or the other of the two
sourcesof illumination. There was no sign of a resultantreaction.
Even if theindividualsporangiophore did not receiveequal illumina-
tion fromthe two openings,if therewere any resultantreaction,it
wouldbe expectedthat the sporangiumwould be foundin a position
betweenthe two lights,dependingon the ratio of theirintensities,
differencesofcomposition,and thelike. Thus, it would be expected
that all of the sporangiophoreswould be located between the two
sources of illumination. Such a conditiondid not obtain in any
case where the two light stimuliwere used, whetheror not they
were the same as faras distributionin the spectrumwas concerned.
The sporangiophorereacted to one or the otherof the two stimuli.
The resultsobtained at least suggest that one stimulus does not
ii8 BOTANICAL GAZETTE [FEBRUARY

affectthe reaction to the other. To this degree the results are

corroborativeof NOLL'S theory. But NOLL'S theory,based on his
work with lightand gravityas stimuli,suggestsa change of geoto-
nus due to the presenceof the light. In workingwiththe two light
stimuli,the reactionto one of the two stimuli to the exclusionof
the othercannot be explainedin this way, althoughhere,as above,
the plant is subjectedto two directiveinfluences.
Where the two simultaneous stimuli were of differentkinds,
gravityand light,NOLL believed that lightmay call forthcertain
changesin the plasma which,directlyor indirectly,cause the reac-
tion. He says that perceptionand reaction may rest on entirely
different characters; but the reaction may be carried out by the
same changes. In work where the two simultaneousstimuli are
of the same character,as in the presentexperiments,the reaction
is less complicated. Thus but two sets of changes of the plasma
need be concernedwith the response.
The above experimentsshow that thereis no sign of a resultant
reaction,but that they can in no way determinewhether,so far as
perceptionis concerned,there is any influenceof one lighton the
other. According to NOLL'S theory,a change of geotonus takes
place, due to stimulationby light; that is, there is a change of
sensibilityto gravity as such due to the perceptionof light. The
question arises whether,withthepresenceof two lightstimuliof the
same kind acting throughthe same time, thereis a corresponding
changeof sensibilitytowardone lightowingto themerepresenceof
a second of a similarkind. Apparentlytheremust be some factor
or factorssomewherein the organizationof the plasma that brings
about a total neglectofone stimulusand a completereactiontoward
anotherof a similarnature,since the reactionis not a resultantone.
It appears thatNOLL'S theoryalone is insufficientto explainentirely
of
the lack resultantreaction to two directiveforces when applied
to an organismas contrastedwiththe resultsobtained by physicists
in workingwith inanimate matter. The question, then, of what
determinesthe reaction toward one light and a lack of response
toward the second is still unsettled,and the explanationmust be
deferredto a timewhen more is knownof the intricatemechanism
and ultimateorganizationof the plasma of the cell.
I9I4] JOLIVETTE-PILOBOLUS i19

In the experimentwith the differentincandescent filament

lamps, as with the solutionsand plates of coloredglass used in the
earlier work (i), Pilobolus fires its sporangia in larger numbers
towardthelightsin whichtheproportionof theblue raysis greatest.
In otherwords,it is more responsiveto actinic rays. The intensi-
ties in the differentwave-lengths,as earlier mentioned,are not
measurable; but the uniformityof responsein favorof the source
containing the greater proportion of actinic rays suggests the
superiorityof the more refrangiblerays over the less refrangible
rays in causing heliotropic curvatures. This question can be
definitelysettled only when the intensitiesof lights of different
colors can be measured.
The energygiven offby the source of lightapparentlydoes not
compare in effectwith the distributionof the same in different por-
tions of the spectrum. In the experimentsusing a 20-watt or
i6-candle-powertungstenlamp and a 32-candle-powercarbon fila-
mentlamp the largemajorityof the sporangiawentto the tungsten,
although its total energywas but half that of the carbon. From
in distributionin the spectrum
this it is apparent that differences
outweigh in effectthe in
differences the total energyof the two
sources.
The set of experimentsusinga i6-candle-powertungstenagainst
a tantalum of twice the numberof candle-powersshowed the dis-
charge to be in favor of the tantalum. At firstglance it appears
that thiscontradictsthe above resultswiththe carbonand tungsten
lamps, and suggeststhat the total energyof the source does play
an importantrole in the results. However, on furtherconsidera-
tion, it must be noted that the total numberof actinic rays in a
tantalumlamp of twice the intensityof the tungstenis probably
greaterthan that in the tungsten. The solution of this point, of
course,is bound up with the question of intensityand composition
of the sources under discussion and cannot be taken with any
degree of finality. With the tungstenlamp of approximatelythe
same intensityas that of the tantalum,the dischargewas in favor
of the tungstenwhich emits a largerproportionof the blue rays.
The differenceas far as distributionin the different wave-lengths
of the energyof the tantalum and tungsten lamps is not so great
I20 BOTANICAL GAZETTE [FEBRUARY

as is the case of that in the carbon and tungstenlamps. This may

account forsome of the differences in the distributionof the spo-
rangia in the two cases, when comparing the tungstenwith the
carbon and with the tantalum.
A comparisonof the carbon and tantalumlamps shows again a
majorityofsporangiadischargedon theopeningbeforethe tantalum
lamp whichcontainedthe largerproportionof blue rays.
In the experimentswith the rays of differentwave-lengths,
although the stimuliare both lightstimuli,thereis a marked dif-
ferencein composition. We have then an extra factor to deal
with; but, as in the experimentswith the two nearly equal light
sources,thereis no sign of a resultantreaction. There is no sign
of a change of aim toward one light owing to the presence of a
second light. With the tendency of the sporangiophoresto dis-
chargetowardthe blue light,however,it is plain that there is no
uniformaim of all the sporangia subjected to the two lights to go
to the lighthaving the largerproportionof the blue rays. Why
does not Pilobolus always dischargetoward the source oflighthav-
ing more of the actinic rays? The differencein the lengthof the
lightray does bringa markedvariationas regardsthe numbersfired
toward the two sources. Still, some are firedtowardthe less favor-
able of the two sources.
The accuracy of aim toward the two lightsmightwell be said
to agreein generalwiththat already foundforthe two lightsources
used in the above experiments. However, there is a noticeable
differencein accuracy of aim toward the differentfilaments,and
that for the most part is in favorof the lightwith the largerpro-
portion of the more refrangiblerays. With the solutions and
glass plates used in the earlierwork (i) therewas a much greater
difference noted. There is a probabilitythat the smallerdifference
may be due to less difference in lightintensity,to a smallerdiffer-
ence in composition,and also that thereis a limit to the accuracy
of response toward any source of stimulation,and that in aiming
at the lightsin use in these experimentsthey reached that limit,
the less effectivelamp being sufficient, or in some cases nearly
so, to bring about as accurate a reaction as is possible to the
plant.
I9I4] JOLIVETTE-PILOBOLUS I2I

The question as to what propertiesof the protoplasmcause it

to be more sensitive to rays of one wave-lengththan to those of
anotherremainsunsolved, and, like that as to why it responds to
one of two stimulito the complete exclusionof the other,it must
await a betterknowledgeof the organizationof the plasma.
LELAND STANFORD JUNIOR UNIVERSITY
CALIFORNIA

LITERATURE CITED
i. ALLEN, R. F., and JOLIVETTE, H., SomelightreactionsofPilobolus. Trans.
Wis. Acad. Sci. I9I2.
2. GUTTENBERG, RITTER VON, Uber das Zusammenwirken von Geotropismus
und Heliotropismus Pflanzentheilen.Jahrb.Wiss. Bot.
in parallelotropen
45:I93-23I.. I907.
3. , Uber das
Zusammenwirken von Geotropismus und Heliotropismus
und die tropistische in reinerund unreinerLuft. Jahrb.
Empfindlichkeit
Wiss. Bot. 47:462-492. I9IO..
4. NOLL, F., Heterogeneinduction. Leipzig. I892.
5. RICHTER, O., Uber das Zusammenwirken von Heliotropismusund Geo-
tropismus. Jahrb.Wiss. Bot. 46:48I-502. I909.