The Spiral-growth of Phycomyces - Wageningen UR E

The Spiral-growth of Phycomyces - Wageningen UR E
KONINKLIJKE AKADEMIE VAN WETENSCHAPPEN
TE AMSTERDAM
The Spiral-growth of Phycomyces
COMMUNICATION N°. 31 OF THE LABORATORY FOR PLANTPHYSIOLOGICAL RESEARCH, W A G E N I N G E N , H O L L A N D .
BY
A. J. P. OORT
Reprinted from: Proceedings Vol. XXXIV, N°. 4, 1931
. f BIBLIOTHEEK^
Botany. — The Spiral-growth of Phycomyces. By A. J. P. OoRT. (Communicated by Prof. A. H. BtAAUW.) (Meded. N°. 31 van het
Laboratorium voor Plantenphysiologisch Onderzoek, Wageningen,
Holland.)
(Communicated at the meeting of March 28, 1931.)
I n t r o d u c t i o n . In one of my experiments on light-growth-responses
in Phycomyces I was struck by the fact that the otherwise quite globular
sporangium showed a small excrescence on one side. The remarkable part
of it was that on examining the sporangium sideways with the aid of a
cathetometer, this excrescence grew gradually larger, next again decreased
in size and finally became quite invisible. After some time, however, the
excrescence appeared on the other side of the sporangium to disappear
from there likewise after a while. After an other interval of time it appeared
on the first side, etc. It was obvious to assume therefore that the sporangium
turned round a vertical axis. Various questions now immediately arose. Do
all sporangia rotate or is it only an abnormality ? Is the rotation connected
with the growth in length or does only the sporangium turn and not the
sporangiophore ? At what rate do they rotate? Before our going into these
questions something should be communicated about the method followed
M e t h o d . To render the rotation visible I used very fine glass needles,
which I glued to the top of the sporangium with minute drops of saliva or
liquid gelatine. Originally I used glass needles of a diameter of ± 60 tu and
a length of about 2 mms. From a calculation it appears, however, that the
weight of a similar glass needle means a not to be neglected increase in
weight of the sporangium. The sporangia namely have an average diameter
of 600 f.i. If we put the specific gravity = 1 (which is certainly not to
much), such a sporangium weighs 4/3 n (0.3) 3 mgrs = 0.11 mgr., while a
glass needle of a length of 2 mms and a diameter of 60 /t, weighs
0.014 mgr., if we put the specific gravity = 2,5.
So the average increase of weight is round 13 %. Afterwards I found in
glasswool a material that is thinner yet, but which is consequently much
more difficult to handle. The average diameter was 35 /t in this case and
the weight of a needle not yet 0.005 mgr., so that the weight of the
sporangium is but increased by 4 to 5 %. Subsequent experiments wers
always made with these lighter needles. A fundamental difference between
the two kinds of weight-increases, however, was not stated.
I measured the rotation by means of a horizontal microscope (fig. 1: a)
via a mirror (b) placed over the sporangium at an angle of 45°. Initially
565
the observations were made in a faint white light, falling in the thermostat
from above. The light reflected by the needle, however, was quite insuffi-
Fig. 1. The apparatus.
cient in certain positions of the needle, so that I rejected this method and
searched for a method of measuring in light, which is send through the
object (just as is done in the case of measuring the growth in length). To
attain this end we must have at our disposal sporangia which viewed from
above grow over the edge of the culture-pot, so that a source of light can
be placed under them. Sporangia meeting with this requirement I obtained
by placing the culture-pots horizontally for some time and next putting
them vertically again. This treatment produces sporangia that are twice
bent (fig. 1: c ) . The pot is placed on a piece of red glass (d) under
which a diminutive electric lamp (e) has been put in a closed space. For
clearness' sake 2 of the 4 reflecting mirrors have been left out in the
drawing. If we subsequently place the culture with its accessories in the
thermostat, we look via the mirror down upon the top of the sporangium
and the needle, with a red field as background. The rotation was measured
by turning the scale of the eyepiece-micrometer in such a way that the
strokes of the scale run parallel with the needle, while the angle was read
directly from a graduated arc by means of a long hand (f) fixed to the
ocular. The size of this graduated arc allowed of a least count of 0.2°. From
a non-rotating (mature) sporangium it was determined how great the
accuracy of adjustment is. The extreme values differed ± 0.2—0.3° from the
average. The accuracy of reading and of adjustment might both be easily
increased by a more accurate scale and a higher magnification, but for this
research the accuracy attained was amply sufficient.
As fig. 1 shows it is possible to measure the growth in length simul-
566
taneously. For this purpose is the cathetometer (g) and the red light (h)
mounted behind the thermostat.
A -j- culture of Phycomyces Blakesleeanus Burgeff (see BURGEFF 1925)
was used, the same species with which BLAAUW and TOLLENAAR worked
under the name of Phycomyces nitens and which was cultivated in the way
indicated by BLAAUW (1914) in pots with moist bread.
To make the sporangia grow in the desired position (twice bent) they
were cultivated in the light. Geotropism by itself does as a rule not suffice
to make the sporangia raise themselves quite from their horizontal position.
Putting on the needles also occurred in white light. Next the sporangia were
placed in the thermostat (described by BLAAUW 1914) at 17.5° in the dark
and I waited for at least an hour before I started measuring. Though after
one hour the light-sensitiveness has not yet increased to the dark-value,
the dark-growth-response is finished by that time and consequently the
growth in length has again become about constant.
R o t a t i o n a g e n e r a l p h e n o m e n o n . From more than 20
experiments in constant temperature and from various observations under
variable conditions it appears that rotation is a general phenomenon shown
by all growing sporangia. (Plate 1). But seldom, and then but for a very
short period the rotation is so slight that it is scarcely measurable and
accordingly there seems to be no rotation.
Before our discussing the measurings of the rotation the question needs
first to be put in what part of the sporangium or of the sporangiophore the
rotation originates.
T h e l o c a l i z a t i o n of t h e r o t a t i o n . Above it was already
observed that the rotation occurs in growing sporangia and therefore it may
obviously be assumed that this rotation occurs in the same zone in which
the growth takes place. The growth is confined to a small zone lying at
± 3^2 to 2—3 mms' distance below the sporangium. (See ERRERA 1884).
The following observations and experiments prove this supposition to be
correct.
1°. Mature sporangia do not rotate.
2°. Sporangiophores which have just formed their sporangium and are
therefore in the stage (stage 3 of ERRERA), in which the growth is quite
stopped, do not rotate. Simultaneously with the beginning of the growth in
length (stage 4 of ERRERA), rotation also commences (Fig. 2).
3°. The rotation shows a reaction after exposure to light. This reaction
proceeds qualitatively and in the main also quantitatively parallel with the
reaction which the growth in length shows after the same exposure. The
graphic representations (figs 3 and 4) demonstrate this parallelism clearly.
Fig. 3 illustrates the response of a sporangiophore growing in faint white
light ( ± 10 M.C. 4-sided) which was exposed to 40.000 M.C.S. 4-sided.
In A the time in minutes has been plotted on the abscis, on the ordinate the
3
567
10-
16»
I8 H
2!0 H
22"
2^H
-2
Fig. 2. Rotation and growth in length during the greater part of stages 3 and 4.
Abscis : time in hours. Observations were made at points of time marked j .
Ordinate :
Rotation in degrees per min. Left-handed rotation negative.
Growth in ,« per min.
s
4-0 MIN.
50
0
IQ
20
30
Fig. 3. Light-response of growth (A) and of rotation (B). Abscis: time in min. Ordinate:
A. growth in /i per min., B. rotation in degrees per min. Continuous exposure to
± 10 M. C. 4-sided. At time 0 (f\ short exposure to + 40.000 M.C.S. 4-sided.
40
568
increase in length in /< per min. At t (time 0) exposure took place.
Y
Observations every 2 minutes. In B. the rotation in degrees per min. of the
same sporangium has been plotted on the ordinate. Here too observations
5ÛMIN
0
10
20
30
Fig. 4. Light-response of growth (A) and rotation (B). Sporangium in the
At time 0 ( T ) exposure to + 1 6 0 M.C.S. 4-sided. Further as in fig.
Ml
darkness
3.
were made every 2 minutes ; it should, however, be noted that the periods
of observation alternate with those concerning the growth in length. One
minute an observation was made through one microscope, the next minute
through the other microscope.
Fig. 4 gives the responses of a sporangiophore growing in the darkness,
which at t (time 0) was exposed to ± 160 M.C.S. 4-sided. A again gives
the increase in length in //.per min. B. the rotation in degrees per min. W e
see that the curves of A and B in both figures especially as regards the
maximum run beautifully parallel. After that the agreement is less perfect,
but it frequently occurs that at this very point of time (so after the maximum) little curvatures appear, which will affect both the measuring of the
growth in length and that of the rotation, that is now in the same sense,
now in the reverse sense.
Even if these first 3 arguments are not absolute evidence, because two
569
simultaneous events need not necessarily happen in the same place, the last
(4th) argument leaves no doubt about it.
4°. If we apply glass needles in and below the growing zone, it
appears that needles beiow the
growing zone do not turn, whereas
needles in the growing zone initially
rotate fairly rapidly, later slower
•
2"30 and finally stop altogether. The
needle has turned out of the rotating zone and also out of the
growing zone, for at the same time
the increase in length of the zone
/">
lying
below the needle has also been
H
reduced to nil.Rotation and growth,
therefore, take place in the same
'• zone. The above has been schematically represented in fig. 5 a, b and
c. In b we see a sporangiophore
H
with three needles, one on the top,
one in the zone of growth and one
H
°
below the zone of growth. In c we
see the same sporangium some time
_
later. The sporangiophore has
u
Fig. 5. Schematical representationofspiral- grown a great deal, while simultagrowth. a. Sporangiophore with zone of neously a rotation of nearly 180°
growth; b. with needles above, inandbelow h a s t a fc e n p l a c e . The lower part of
the zone of growth ; c. the same sporangio
phore some time after ; d. the path of a point
above the zone of growth, when we think
the cylinder of the sporangiophore spread
out in one plane and when we look from
t he outside.
the zone of growth (below the
middle needle) has grown less,
while the rotation is also slighter
(nearly 90°). The lower needle
shows that belowthe zoneof growth
no rotation takes place. Fig. 5d shows the path described by a point on the
wall above the zone of growth when we think the cylinder of the sporangiophore spread out in one plane and when we regard it from the outside.
S pira 1- grow th. From the conclusion : "growth and rotation always
go together and are confined to the same zone", it follows that the actual
direction of growth does not coincide with the longitudinal direction of the
sporangiophore, but forms a definite angle with it. The direction of growth,
therefore, runs spirally round the cell. Accordingly we have called the
phenomenon s p i r a l - g r o w t h .
W e will now further analyse this spiral-growth. W h e n measuring the
rotation, properly speaking we measure the width-component of growth ')
l
) One might be inclined to speak of g r o w t h in width, but we had better avoid this
term. For the growth (increase in volume of cell-wall) exclusively takes place in a longi-
570
and when we know the diameter of the sporangiophore (which was
measured at the end of each experiment), we can express this widthcomponent in fj, per min. instead of in degrees. From the ratio of the lengthand the width-component, which were measured at the same time, the angle
can then be computed which the main direction of growth forms with the
longitudinal axis.
In table 1 the results of more than 20 experiments have been resumed.
W h e n first we observe the direction of the rotation, it strikes us that it is
right-handed 1 ) in 17 cases. In the other 5 cases the rotation is partly
right-handed and partly left-handed.. (In the table and also in fig. 2 lefthanded rotation has been marked — ) . Sporangia rotating exclusively lefthanded or not rotating at all were not found. Accordingly the spiral can
change its direction and pass from right-handed to left-handed of from
left-handed to right-handed. But it is indeed obvious that the right-handed
spiral is being preferred. It is not yet possible to explain the reason why
one sporangiophore behaves differently from another. Therefore I will here
only indicate some points that may probably throw light on this question. If
beside the rotation we also consider the growth in length, it strikes us that
the sporangiophores with exclusively right-handed rotation as a rule grow
most rapidly (30—60 /t per min.), while most sporangia with left-handed
and right-handed rotation grow much slower (10—20 fj, per min.) and
consequently we get the impression that there is something wrong with the
growth in these sporangia. W e should then have to consider the sporangiophores with alternate left-handed and right-handed rotation as abnormal
cases.
This assumption is strongly supported by the process of the growth of
sporangium N°. 1 from table 1. Initially the growth was normal and the
rotation right-handed. Both, however, decreased till at a certain moment
both stopped altogether. I thought the sporangiophore had arrived at the
end of the grand period, but after a short time the growth recommenced
and so did the rotation, but it was now left-handed. Growth and rotation,
however, were slow (respectively 10—15 fj, per min.and 1°.5—3°permin.).
In normal cases no stagnation of growth occurs in the grand period ; it is,
therefore, not strange to regard left-handed rotation in this case as abnormal. The growth of the sporangiophores nos .8, 14, and 19 is now
comparatively slow, only N°. 13 with an average growth of 47 fi per min.
forms an exception. In sporangium N°. 19, of which the greater part of the
tudinal direction. By growth in width one should rather understand an increase of circumference of the sporangiophore. This would result in a widening of the sporangiophore
towards the top, which occurs for instance in Pilobolus and some varieties of Phycomyces
Blakesleeanus. This growth in width is here out of the question.
') W e call a spiral right-handed, when the observer who considers it as a winding
stair-case and ascends it, has the right hand at the inside. W h e n we look down on the
sporangium from above we see in the case of a right-handed spiral the needles on the
top of the sporangium move like the hands of a clock.
7
571
T A B L E 1.
N".
exper.
Duration
exper.
in hours
Average
growth
in length
in ii min.
Direction
of
rotation
1*
3
—
R-L
2*
I'/j
55
R
Average
rotation
in
degr. min.
Diameter
sporangiophore in/i
Average
widthcomponent in
fi min.
Average
z«
in degr.
3*
1'A
—
R
3.6
2.3-4.7
4
2
46
R
4.5
2.4-67
—
—
—
—
5
37
R
2.1
1.2-3.1
90
6*
3'/2
4
46
R
1.5
0.2-3.7
108
1.4
1.8
7
4
33
R
2.6
1.4-3.3
120
2.7
4.7
8*
4'/2
15
L-R
9
172
40
R
10*
2'/2
1
18
40
12
1
13*
3
M*
6
11
15
—
Min. and max.
rotation
in degr. min.
—4.5-0-9.0
6.7
5.0-8.2
—
—
—
—
—
1.6
—
—
—
—
—
2.5
—
—5.3-0-7.0
64
7.1
5.8-10.7
156
R
1.1
0.0-2.0
R
2.9
2.2-5.1
120
3.0
4.3
26
R
3.5
4.2
9.2
47
R-L-R
13
L-R
—
—
—
—
2.3-4.8
136
—1.3-0-1.8
162
—5.6-0-1.1
97
—
—
9.7
—
—
—
—
13.5
—
—
—
—
R
2.7
16
2
63
R
4.8
3.2-8.5
83
3.5
3.2
17
6
43
R
2.9
1.0-4.5
102
2.6
3.5
12
R
1.6
0.9-2.5
110
1.5
7.1
18
19*
72
372
16
21
R-L
72
3
R
0.5
20
—
—1.9-0-2.0
—
—
—.
—
—
—
—
21
5
54
R
6.9
3.8-9.5
98
5.9
6.2
22
4
36
R
4.4
3.1-5.3
130
5.0
7.9
113.9
3.74
5.81
Average*
39.1
3.65
*) Observations. Left-handed rotation negative.
No. 1. Average growth during l'/2 hours 43 ,n, next
stopping of growth followed by slow growth of
12 (i min. and slow rotation L. (see also text).
Continuous light 10 M.C.
No. 10. Rotation during short time practically 0.
No. 2. Continuous light 10 M.C.
No. 14. During 2'/2 hours L., then 3'/2 hours R.
Average growth in length slight!
No. 3. Measured from photographs of plate 1.
Temp. 16°.
No. 6. Rotation short time almost 0.
No. 13. During V/2hours R., then 1 hour L., next
V2 hour R.
No. 19. Average growth computed after last 9 hours.
During 9 hours R., then 5'/2 hours L. See fig. 2
and text.
No. 8. During 1i/i hours L., then 3Y4hours R. Growth
in length slight.
T h e average was computed from 11 experiments; rotation exclusively R. viz. Nos. 5, 6, 7, 9, 11,
12, 16, 17, 18, 21 and 22.
572
grand period was measured, the rotation was first right-handed for nearly 9
hours, next for 5x/i hours left-handed (see fig. 2), but here too the average
growth is subnormal and the final length
of the sporangiophore amounted only
to 3.5 cm, 10 cms being normal. The
culture employed was comparatively old,
so that a deficit of nutrient substances
or water may be the cause of this
abnormal behaviour.
These are the arguments supporting
the supposition that the left-handed
spiral probably does not occur but in
abnormal cases.
Regarding fig. 2 more closely, we see
that in the positive part of the curve
Fig. 6. a. Possible explanation of alterconsiderable oscillations occur in the rate
nating R and L rotation by oscillations
of rotation. It also appears from the
in the direction of growth round an
other
experiments (table 1) that the rate
average direction forming a definite angle
of
rotation
is not constant. These oscilla(/_ tt) with the longitudinal axis. b. The
direction of growth of a R—L—R rotations proceed gradually and the maxima
ting sporangiophore drawn on the wall,
and minima are at least two hours,
viewed from the outside. Both schemafrequently much farther removed from
tically.
each other. Besides long periods occur
in which oscillations fail to appear. From these oscillations the left-handed
rotation can now also be explained. W e must then assume that the average
main direction of growth forms a definite small positive angle (a) according to a right-handed spiral with the longitudinal axis, but that the oscillations can be so large that temporarily negative angles can also occur, that
is left-handed rotation can take place. This has been schematically
represented in fig. 6 : a. A good explanation of the rotation of sporangiophore N°. 13 (schematically represented in fig. 6b) might be given in
this way.
This also applies to the sporangiophores Nos 6 and 10, the rotation of
which is indeed always right-handed, but which for a short period hardly
rotate at all. If the oscillation had been a little stronger a left-handed spiral
would have occurred for some time. But if we now regard fig. 2 it appears
that the above does not give a satisfactory explanation for all cases, because
in this case the left-handed rotation continues too long for that. Growth and
rotation, therefore, belong together, but as the steepness of the spiral in
one and the same cell varies in the course of the hours and also among the
different cells, there is no fixed relation between the velocity of growth in
length and that of rotation. The rotation, however, does increase and
decrease along with the growth, just as in the case of the response to light.
A t w h a t r a t e d o e s t h e r o t a t i o n t a k e p l a c e ? The table
may give us a notion of this, but for the reader it is easier to form an idea
10
573
of the rate of the spiral-growth with the aid of some round averages. For
these averages I only made use of the sporangia with right-handed rotation.
The average growth amounts to 40 /.t per min., the average rotation to 4°
per min. With a circumference of the sporangiophore of 360 JX, the component of width then amounts to 4 ju, per min. The angle the main direction
of growth forms with the longitudinal axis {/(i) is then ± 6°. Such a
sporangium will have made a complete revolution in 1Yi hours, the increase
in length in that period amounting to 3.6 mms. If we now put the increase
in length during the grand period at 8 cms, the end of the cell with the
sporangium, there fore, turns more than 20 X round its axis during
that period.
S p i r a l - g r o w t h a n d s p i r a l - s t r u c t u r e . In the literature
on the growth and structure of the cell-wall cells are repeatedly mentioned
showing a spiral-structure ; but no quantitative data about spiral-growth
are found. The word spiral-growth however is used several times. But in
most cases the word is used for organs, which are composed out of more
cells and so it is dubious if we may compare these cases directly with the
spiral-growth of Phycomyces. Yet I have to mention the investigations of
PRINGSHEIM und LANGER (1924) on Bacillus mycoides and those of
KLUYVER und VAN N I E L (1926) on Bacillus funicularis, especially where
the autors speak about "Zopfbildung". It is hardly possible to explain this
"Zopfbildung" otherwise than through the spiral-growth of the bacteriathreads. Also I have to mention an observation of BURGEFF (1914 p. 279)
on Phycomyces nitens var. piloboloides which I quote here : "Es entsteht
unter einer Dehnung der Membran (des Sporangiumträgers) eine blasige
Anschwellung unter dem Sporangium. Während dieser Dehnung erleidet
der blasig werdende Teil des Trägers eine Torsion von links nach rechts,
die an den in ihm spiralig ausgespannten Plasmafäden sichtbar wird." It
is not quite clear if BURGEFF really observed a torsion, but according to
this observation about the direction of the plasma-threads it is very
probable that this variety also will show spiral-growth.
As regards spiral-structure we find it for instance very common in fibres.
1926 and 1927). In many cases this structure can already be recognized by striation or by the direction of bordered pits, but with the aid of
the polarization-microscope the structural direction can always be determined
more exactly. For according to NÄGELl's micellar theory all cellwalls have
been constructed from submicroscopical crystalline particles, the so-called
micellae. Whether these micellae consist of one single large molecule or of
a number of molecules, is not yet known, because the molecular weight of
substances as cellulose and chitin is not known. The particles are, however,
crystalline which appears from the fact that they show double refraction
and yield X-ray interferences.
In many cases the crystals would show the shape of rods x), of whicii
(FREY
') See also BAAS—BECKING and WAYNE GALLIHER.
A. J. P . ' P O R T :
THE SPIRAL-GROWTH OF PHYCOMYCES.
Photographs of the rotation, taken in red light ; magnification almost 5)*(. Time of
exposure ','2 minute: exposures every 15 min. Temp. 16°.
Proceedings Royal Acad. Amsterdam, Vol. X X X I V , 1931.
PLATE I
574
11
the longitudinal direction coincides with the greatest coefficient of refraction, that is with the axis of the (approximately uniaxial) crystal. The
rods (micellae) now always point more or less in a certain direction and
the direction coincides with the direction of a striation or of thickening
ridges. Accordingly the structural direction can be determined and as
already stated it appears that a spiral-structure frequently occurs. Within
this scope I cannot enter into this question further. A good survey of
various structures is found in VAN ITERSON (1927) and FREY (1930). Here
we also find additional literature.
I have dwelled upon the micellar theory, because it may obviously be
assumed that spiral-growth and spiral-structure are connected and that
direction of growth and of structure coincide. In this case these cells of
Phycomyces would be an example in which the spiral-structure is
manifested in the growth, — owing to their being free cells. Theoretically, however, direction of growth and structure need not coincide. It
may for instance be imagined that the direction of growth coincides with
the longitudinal axis, while the particles stand obliquely with respect to
this axis. Compare for instance a herring-bone clinker-pavement. This is
a case in which the parts (clinkers) are orientated obliquely with respect
to the main direction in which the "growth" took place during the formation.
In the same way we might picture to ourselves a cell with spiral growth,
the parts of which are orientated in the longitudinal direction.
In connection with this I also wish to point out the relation found by
different investigators (CRÜGER 1855, DiPPEL 1868, STRASBURGER 1882)
between the protoplasmic streaming and the structure. Here too further
researches on Phycomyces will be capable of throwing light on the question
whether the streaming of the protoplasm determines the direction of growth.
As to the alternate right-handed and left-handed rotation of Phycomyces
it is interesting to point to the "reversions" found in the structural direction
of cotton-hair (see on this subject VAN ITERSON 1927).
Moreover I also wish to point out here that the phenomenon of spiralgrowth in Phycomyces appears in its simplest form. Organs constituted
of more cells often show a spiral-structure as for instance the stalks of
creepers, stems of trees. It is not impossible that the spiral-growth of one
single cell, as shown by Phycomyces, is the elementary phenomenon from
which these more complicated phenomena of growth might be explained.
Wageningen, Maart 1931.
Laboratory [or Plantphysiological
Research.
LITERATURE CITED.
BAAS—BECKING, L. G. M. en GALLIHER, E. WAYNE: Wall structure and mineralization
in Coralline Algae. Journ. Phys. Chem. 35. 467. 1931.
BLAAUW, A. H.: Die Perzeption des Lichtes. Ree. Trav. bot. néerl. 5. 209. 1909.
12
575
BLAAUW, A. H.: Licht und Wachstum I. Zeitschr. f. Bot. 6. 641. 1914.
Licht und Wachstum III. Med. Landbouwhoogesch. Wageningen. 15.
91. 1918.
en V. HEYNINGEN, W . : T h e radium-growth-response of one cell. Proc.
Royal Acad. Sc. Amsterdam. 28. 403. 1925.
BOAS, F.: Lieber Pilzkolonien mit spiraligem Wachstum. Centralbl. f. Bakteriol. Abt. 2.
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