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VOLUME LVII NUMBER 2
T H E
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
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;
FIG. I2
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
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 ..". ... ...
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
~ inIt lo -O 0O% H N in l
. . . . .. .. .. .
. . .
. . . . . . . . . . . . . . .
. . . w. . . . .
ZN~~~~~~~
.. . . . . . . *.*.
... H . ...
. . . ',. . . . . . .
.
0 _ _ . _ _ _ _.__C,_0
_ _ . _ *_ *_ _ _ _
.* N
PA
Nd . H % . . . .
I0
0 . . ,
vZ . . . . . . . . . . . . o. c .
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
Total
Total tE
H Hd
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.. ... ... ... ...
Total
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
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.