Trans. Br. mycoI. Soc. Vol. 58.

Plate 20
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Vol. 58, Part 2 April 1972
Trans. Br. mycol. Soc. 58 (2), 179-195 (1972)
Printedin Great Britain

PRESIDENTIAL ADDRESS
SPHAEROBOLUS: THE STORY OF A FUNGUS

By C. T. INGOLD

(With Portrait and Plates 21 and 22 and 12 Text-figures)

It would not be easy to imagine a wider subject than 'The Advance of

Mycology'. That was the title of my address to the International Myco-
logical Congress. Today I am, perhaps, going to the other extreme in
talking about a single fungus. Nevertheless, I am in the good company of
a former President, Miss Blackwell, who in 1942, devoted her address to
Phytophihora cactorum.
Long ago it was suggested that gasteromycetes can be interpreted as a
series of remarkable experiments in dispersal. It was postulated that the
development of these experiments became necessary with the loss of the
hymenomycete equipment of spore liberation; equipment based on a
spore gun, the active basidium, with a range rarely exceeding 0'2 mm.
These experiments (Fig. I) included fruit bodies of the puff-ball
type; the phalloids adapted in relation to insect dispersal; subterranean
forms apparently relying on mycophagous rodents to spread their spores;
and the beautiful bird's nest fungi, Cyathus and Crucibulum, depending
essentially on large rain-drops for the splash-dispersal of their peridiola.
There has even been developed, and to me especially this seems a little
miracle, a submerged aquatic gasteromycete, Nia vibrissa, found growing
on wood in the sea (Doguet, 1967). This has basidiospores fantastically
like the conidia of the aquatic hyphomycetes, which I know so well, with
their long divergent arms (Fig. 2).
However, it will be generally agreed that of all the gasteromycete
experiments Sphaerobolus is the most spectacular. As in other gastero-
mycetes this extraordinary fungus no longer has short-range basidial guns,
but has developed secondarily a refined catapult mechanism of discharge
with a range unequalled by any other fungus.
Sphaerobolus has attracted attention for nearly two and a half centuries.
Micheli's original figure (PI. 2 I) in Nova Plantarum Genera published in 1729
illustrated its essential structure and activity (Micheli, 1729). A full
summary of the histological and biological work on Sphaerobolus was
beautifully presented by Buller in 1933 in Vol. V of his Researches onfungi
in nearly 100 pages of exciting reading. He drew extensively upon a major
paper by Walker (1927) and added many original observations.
The taxonomy of the genus needs critical study, but probably most
collections from all over the world can be referred to the type species:
S. stellatus Tode ex Pers. Certainly all British material I have seen appears
to belong to a single species. However, two isolates from Mrica and one
Vol. 58, Part I. was issued 28 February 1972
12 MYC 58
180 Transactions British Mycological Society

Fig. I. Cartoon illustrating various types of dispersal in gasteromycetes. Scales

very different for the different fungi.

Fig. 2. Nia vibrissa. Basidiospores from sporophores on submerged wood in Irish Sea.
 Presidential address. C. T. Ingold 181
from Pakistan which I have grown seem so different that they must surely
be placed in another species (Ingold, 1971). As compared with British
isolates they are much larger and the limiting wall of the black glebal-mass
is structurally different from the brown one of other isolates. Again the
glebal-mass has a distinct 'cap' and within, in addition to basidiospores
and germinating gemmae, are some spherical 'cystidia', capable of
belated germination. These seem to be lacking in the smaller species.
Two isolates of the smaller species from warmer countries have been
studied. One of these is from Ife in Nigeria and the other from California.
They agree essentially with isolates from Britain not only morphologically
but also in growth rate and in their temperature characteristics (Alloway
& Ingold, 1971).
My talk today is based largely on my concept of S. stellatus as represented
by our Isolate A. This has been used for experimental purposes in my
laboratory continuously for a dozen or more years.
The consideration of any life-cycle may start at any point and we may,
therefore, begin with the discharged projectile. Within the brown glebal
membrane of disorganized hyphae are tens of thousands of rather thick-
walled, uninucleate basidiospores. There are also hundreds of larger, thin-
walled, dikaryotic gemmae poised for immediate growth (PI. 22). The
basidiospores, however, seem to need special conditions for germination,
conditions which may, perhaps, be realized in nature during passage
through the alimentary canal of a herbivore. In the laboratory it appears
that they have been successfully germinated after treatment with pepsin.
My own efforts in this direction have, however, always failed. In our
isolates of S. stellatus there are also extra-glebal gemmae, quite distinct in
form from those of the gleba, on the surface of the projectile. When this
strikes an object, some of these are spattered into the air thus providing a
possible means of airborne dispersal (Ingold, 1971). They germinate
freely. The isolate from Ife in Nigeria has unusually small sporophores
and the glebal-mass is only about 0'5 mm in diameter. Further, the mem-
brane around the gleba has little pigment and is, therefore, rather trans-
parent. Because of these features when a discharged glebal-mass caught on
a slide is gently flattened under a coverslip, the extra-global gemmae can
be seen in situ on the surface (Fig. 3).
We almost always collect Sphaerobolus on our autumn forays and the
great majority of these collections are from very rotten wood. I remember,
however, that we found the fungus with considerable frequency on old
cow-dung pads during the Hereford Foray of 1951. As an interesting aside
it may be mentioned that when the artist-naturalist James Sowerby
investigated wood-rot in the man-of-war Queen Charlotte in 1812, S.
stallatus was one of the species he figured from 'the Dock-yards and Ships
of Great Britain from Deptford to Plymouth'. I make this an excuse for
drawing the attention of my fellow mycologists again to Dr Ramsbottom's
story of dry-rot in ships, surely one of the most joyous chapters in the
literature of Mycology (Rams bottom, 1941).
Sphaerobolus seems ideal for its double ecological status as a lignicolous
and as a coprophilous fungus. If the projectile alights on dead wood in a
suitable state, this can immediately be colonized through the activity of
I2-2
182 Transactions British Mycological Society

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Fig. 3. Sphaerobolus stellatus. Nigerian isolate from Ife. (A) Flattened glebal-mass on lid
of petri-dish. (B) Contents of gleba consisting of basidiospores and gemmae. (C)
Surface view of indicated region showing extra-glebal gemmae in situ.

the gemmae within the glebal-mass. If, however, it comes to rest on grass,
it can bide its time there firmly stuck. If this herbage is eaten by a cow the
dormant basidiospores are, apparently, ready, following stimulation in the
alimentary canal, to germinate in the dung. It is a pity, however, that
experimental study of this aspect of dispersal in Sphaerobolus appears to be
lacking.
S. stellatus has the basic characteristics of such specialized dung fungi
as Pilobolus, Dasyobolus and Podospora (Fig. 4). All four genera have certain
features in common: the projectile is a relatively large spore-mass;
it is thrown to a considerable distance; a phototropic response of the fungus
determines the direction of projection; most of the spores in the spore-
mass do not germinate without special treatment; the spore-mass being
sticky becomes firmly stuck following impaction; and the protoplasm of the
spores is screened from light either by melanic pigment in the spore-walls
or by a dark membrane covering the spore-mass.
In the laboratory the best starting-point for a culture is a freshly dis-
charged glebal-mass. With this procedure our Isolate A has shown no
diminution in fruiting capacity during its many years of extensive use in
my laboratory. Incidentally it is worth noting that Sphaerobolus can be
stored as discharged glebal-masses, preferably in a 'fridge'. It has been
found that even after 12 years, dried specimens can revive and provide
cultures.
 Presidential address. C. T. Ingold

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;1f20emyl
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Fig. 4. Parallel adaptation in coprophilous fungi. (A) Podospora curoula, x 40; (B)
Dasyobolus immersus, x 25; (C) Pilobolus kleinii, x 10; (D) Sphaerobolus stellatus, x 8.

The fungus grows on a wide range of standard media, but for sporulation
nothing has been tried that in any way approaches oatmeal agar. Provided
the illumination is satisfactory and the temperature not too high, fruiting
occurs abundantly. As with a number of higher fungi, an insoluble carbo-
hydrate that only slowly becomes available is, apparently, conducive to
sporophore production. This is probably why oatmeal is such a satis-
factory substratum. If the medium is re-enforced by sucrose at a level of
I % fruiting is greatly reduced, and if that level is increased to 2 % no
sporophores are formed (Fig. SB). It is to be noted that the number of
glebal-masses discharged is an objective measure of fruiting, since dis-
charge is clearly recognizable as the final stage of maturation.
For sporophore production adequate illumination is essential. For
Isolate A this must exceed 100 lux in terms of light from 'daylight'
fluorescent tubes. There is proportionate increase in fruiting with greater
intensity up to a level of 1000 lux, but beyond this, increase has little effect
Fig. SC). However, even if the light is bright enough, the tempera-
ture must also be right. For Isolate A sporophore production has an
optimum around IS °C and almost ceases at 2So which is about the
optimum for extension growth (Fig. SA).
Sphaerobolus fruits well in continuous light and equally well with a daily
regime of 12 h light and 12 h of darkness. No dark period, following
the necessary light one, is needed to complete development such as
Manachere (1970) has reported for Coprinus congregatus.
Although our Isolate A behaves in a satisfactory manner if cultures are
grown from the start in light, greater uniformity of fruiting in parallel
cultures is obtained if they are grown for a fortnight in darkness at 18-20°
184 Transactions British Mycological Society
Temperature A
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Fig. 5. Sphaerabolus stellatus (Isolate A). Features of general nutrition. (A) Inter-
rupted line, temperature and colony radius after 35 days; continuous line, mean
number of glebal-masses discharged from parallel groups of Petri-dish cultures at
various temperatures, after illumination (1000 lux, 'daylight' fluorescent) for 38 days
following initial 2 weeks in darkness at 20°. After Alasoadura (1963). (B) Number of
glebal-masses discharged from samples each of six Petri-dish cultures on oatmeal agar
with and without sucrose supplementation; cultures initially in darkness for 21 days and
then in light (1000 lux, 'daylight' fluorescent) for 44 days, temperature 20°. After
Nawaz (1966). (C) Glebal-mass discharge and light intensity at 20°, cultures 2 weeks in
dark, then in continuous light (' daylight' fluorescent) of indicated intensities for 28 days.
After Alasoadura (1963).

before being continuously illuminated. This closely parallels Manachere's

experience with C. congregatus.
In our Isolate A following illumination of a culture previously in dark-
ness, sporophore initials become visible in a few days' time, and some 15
days later glebal-masses begin to be discharged. Thereafter, under standard
conditions of 20° and 1000 lux, discharge from a Petri-dish culture con-
tinues for several months provided there is sufficient depth of medium and
undue drying is avoided. Further, no general decline in fruiting occurs.
There is, however, a regular periodicity of discharge (Fig. 6C) with
10-12 days between maxima (Alasoadura, 1963; Ingold & Peach, 1970).
There is good experimental evidence that this periodicity is due to young
sporophores inhibiting the development of still earlier primordia. These
apparently become arrested and do not resume growth until the first
crop has matured and discharged (Ingold & Nawaz, Ig67c).
The periodic discharge from a fruiting culture of Sphaerobolus closely
 Presidential address. C. T. Ingold
A
Coprinus

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10 20 30 40
Fig. 6. Flushing in higher Basidiomycetes. (A) Coprinus congregatus: daily per-
centage of culture tubes (22 x 2·2 cm) with new ripe sporophores. Time scale shows
days after start of 12 h darkr r e h light daily regime. Before zero time cultures 10 days
in dark following inoculations. Temperature 24°; light 100 lux, medium: malt agar.
Based on Manachere (1970). (B) Agaricus bisporus: sporophores produced on a mushroom
bed. B I: interrupted lines show number of 'buttons' (0·5--0·75 in. diam.) present over
indicated periods; solid lines show number of mushrooms picked over indicated
periods at 'cup' stage. B2: as above, but mushroom not picked; period of 'buttons'
shown as in B I. Based on Cooke & Flegg (1965). (C) Sphaerobolus stellatus, number of
glebal-masses discharged daily (from the start of discharge) from a Petri-dish culture
on oatmeal agar. Light continuous 1000 lux; temperature 20°. After Alasoadura (1963) .
186 Transactions British Mycological Society
parallels the periodic 'flushing' of sporophores on a mushroom bed. The
basic cause seems to be essentially the same (Cooke & Flegg, 1965). In a
mushroom bed, if the fruit bodies are not picked, the interval between
flushes is about 20 days. If, however, each flush is harvested at the' cup'
stage (i.e. when the partial veil is still unbroken), the interval between
flushes is reduced to 6 days, and the overall production is greatly enhanced
(Fig. 6B). In a Petri-dish culture of Sphaerobolus a flush is automati-
cally harvested by the discharge of the glebal-masses on to the lid. The
exhaustion of each crop of sporophores gives the' all clear' for the next set
of primordia to resume development.
In Manachere's work on Coprinus congregatus, successive crops of new
sporophores also occurred in his batches of test-tube (22 x 2'2 ern) cultures.
Under a standard daily regime (12 h light: 12 h darkness) at 24° crops
appeared at 5-day intervals (Fig. 6A). Here again the explanation
seems to be the same as for Sphaerobolus (Manachere, 1970).
The light factor in the fruiting of Sphaerobolus has been the subject
of considerable study both by Friederichsen & Engel (1957, 1960) in
Germany and by myself and my students at Birkbeck. At 20° about 2
weeks of illumination are required for the complete development of sporo-
phores in a culture previously kept in darkness. Light is absolutely neces-
sary both for their initiation and for their early development. However, if,
after only a week in the light, cultures are again placed in the dark, some
sporophores mature and discharge their glebal-masses, although there is
considerable delay. During most of the second half of the developmental
period light is still a potent factor in maturation. However, in the final
phase, including opening of the sporophore, lasting just over 24 h at 20°
and culminating in glebal-mass discharge, light seems to play no part.
Thus when active, illuminated cultures are transferred to darkness, dis-
charge continues undiminished for a further 24 h, but in the day after that
it is reduced to nothing. Nevertheless, after a few more days in sustained
darkness there is a subsequent slow ripening of sporophores, since light is
not an absolute necessity for the further development of those that have
already completed their first week of illuminated life (Fig. 7B).
At 20° under a regime of I 2 h: light 12 h darkness each day there is a peak
of discharge in every light period: each burst of activity triggered offby light
treatment a day before (Fig. 7A). In the same way each dark-period
minimum relates to an inhibition by darkness experienced 24 h earlier.
If a regime of 12 h light: 36 h darkness is used, discharge, as would be
expected, occurs in the middle of the dark periods (Fig. 7 C). The
response to these two regimes reflects the fact that at 20° the final 24 h
period of development is unaffected by light. At lower temperatures, how-
ever, the duration of this light-insensitive stage is increased. At 10° it is
lengthened by 50 %, so that in a daily regime of 10 h light: 14 h darkness
at that temperature, discharge is almost limited to the dark periods*
(Ingold & Nawaz, 1967c).
So far as the visible spectrum is concerned it has been shown (Alaso-
adura, 1963) that, for overall sporophore development, it is only light of
* No doubt this would also be the case with 12 h light: 12 h darkness each day, but
data from such an experiment are not available.
 Presidential address. C. T. Ingold
A

0
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Days
Fig. 7. Sphaerobolus stellatus: ligh t and glebal-mass discharge. (A) H ourly rat e of
discharge from cul tur es at 20-25° during a period of 4 da ys with the indicated light
regime. After Friederichsen & En gel (1960 ). (B) Daily ra te of discharge from four dis-
cha rging cultures (a t 20°) previously in continuous light an d now transferred to dark-
ness. After I ngold & Nawaz (1967). (C) Rate of dischar ge (per 12 h period ) from four
discharging cultures (at 20°) previousl y in continuous light but from zero time of graph
given th e ind icated light-dark regime. After I ngold & Nawa z (1967). In all figur es the
periods of darkness ar e stippled.
188 Transactions British Mycological Society
shorter wavelength, below 500 nm, that is effective. In this Sphaerobolus
agrees with the great majority of fungi. Nevertheless, in the later stages
of maturation, when light still has a profound influence, a dramatic change
occurs. At around the half-way stage, about a week before maturity,
ripening begins to be hastened by light of longer wavelength, namely
orange and red light in the range 580-720 nm; and at the same time light
of shorter wavelength (e.g. blue light around 460 nm) begins to have a
delaying effect (Ingold & Nawaz, 1967b).
Orange-sensitivity has been studied during the penultimate few days of
development before the final 24 h when light no longer exerts an influence.
The interesting fact emerges that, during this period there is a marked
antagonism between orange and blue light (Ingold, 1969).
A large number of experiments have been conducted in which dis-
charging illuminated cultures were transferred to darkness which was then
briefly interrupted a day later by 1-2 h oflight at a low intensity (100 lux).
In these experiments the daily rate of glebal-mass discharge was recorded
for several days following the initial transfer to darkness. In the period
24-48 h after the interruption with light there was considerable discharge
where orange (585 nm) or red light was used, but virtually none when the
light was blue or green (Fig. 8A). In experiments where the orange
treatment was immediately followed by blue, no discharge occurred, but
where this treatment was reversed the resulting discharge was abundant
(Fig.8B). In other words, the stimulatory effect of orange light can be
reversed by blue. When, however, treatment involves the simultaneous
incidence of orange and blue light, there is no inhibition.
It is difficult to deal adequately with this matter in a general address.
Those interested must read the original papers. Suffice it to say that a
rather similar system has been reported in connexion with the sporulation
of Helminthosporium oryrae (Honda, Sakamoto & ada, 1968), but different
wavebands of light are involved. Near-ultraviolet light (centred around
330 nm) stimulates subsequent sporulation in the dark, but blue (centred
around 420 nm) has a depressing effect. If the ultraviolet treatment is
followed immediately by a brief exposure to blue, the stimulatory effect of
the former is negatived. With short alternating treatments using the two
types of light, there is stimulation if the alternation ends in ultraviolet
light, but inhibition if it ends in blue. Parallel experiments in Sphaerobolus
involving 15 min. alternations of low-intensity orange and blue light give
closely similar results (Ingold & Peach, 1970).
The systems in Sphaerobolus and in Helminthosporium beg comparison with
the red and far-red reversible system in green plants in which phyto-
chrome is implicated.
So far we have been considering light as a major factor in the develop-
ment of sporophores. It is also concerned with their orientation. Positive
phototropism of the fruit bodies was recognized by Walker (1927), by
Buller (1933) and more recently has been studied by Nawaz (1967) in my
laboratory. Both Walker and Buller thought that directional adjustment
to light occurred only in young sporophores, but Nawaz has shown that
even during the final 24 h, when light has ceased to influence the develop-
mental process, the sporophore can respond to a new direction of illumi-
 Presidential address. C. T. Ingold
I II
400 ....,.,.,,-- -,------,- - - ...,.-----, 400 , - -- - - -.........---...,.---,

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Text-fig. 8. Sphaerobolus stellatus. Isolate A. (I) Daily rate of glebal-mass discharge
over 4-day period following transfer from light to continuous darkness (stippled)
interrupted, at times indicated by arrows, by I h orange (Y I), I h red (R I), I h blue
(BI) or I h green (GI) light. Twelve Petri-dish cultures (5 em diam.) in each treat-
ment. (II) Daily rate of glebal-mass discharge over 4-day period following transfer
from light to continuous darkness interrupted, at times indicated by arrows, by I h
orange (Y I ): I h blue (B I); I h orange followed by I h blue (Y I B I ) ; and the reverse
(BI YI). Twelve Petri-dish cultures in each treatment. Temperature 20°; light
intensity for all colours 100 lux. After Ingold (1969).

nation in such a way that the glebal-mass tends to be shot towards the
light.
The early phototropism of the developing sporophore is, apparently,
associated with its longitudinal axis (for it is not quite spherical) becoming
parallel with the incident light. However, the later response to light appears
to be related, not to adjustment ofthis axis, but rather to the determination
of the exact locus in the apex around which stellate opening of the sporo-
phore is to occur. It seems that this affects the final direction of discharge.
It should be said in passing that, although in isolates clearly referable
to S. stellatus phototropic response is a very striking feature, in the larger
species of Sphaerobolus, typified by our South African isolate, phototropic
response is relatively weak.
It has been found (Ingold, 1969) that in this final response to light, which
affects the direction of discharge, the effective light is in the blue region of
the spectrum and orange light has no influence, in spite of the fact that the
reverse is true of the last light-sensitive stage of actual development.
In any organism responding to light there is the problem of the nature
of the chemical photoreceptor. It is to be remembered that, so far as the
visible spectrum is concerned, the great majority of light-sensitive fungi,
190 Transactions British Mycological Society
respond only to blue light. Hence a photoreceptor absorbing in the
spectral region 420-500 nm, and therefore likely to be yellow or orange, is
indicated. There are rival views. Some favour coloured carotenoids; others
look to a flavin, especially riboflavin.
Like so many phototropic fungi (e.g. Phycomyces, Pilobolus and Sordaria),
Sphaerobolus is rich in carotenoids (Schneider, 1964). A methanol extract
of sporophores gives the characteristic absorption spectrum with three
humps in the blue region (Alloway & Ingold, 1971). These pigments might,
therefore, be the essential photoreceptors in connexion both with the over-
all sensitivity of the developmental process to blue light and with the
phototropic response. However, the active receptor for the necessary photo-
chemical reaction might still be riboflavin or something like it. Coloured
carotenoids and riboflavin have absorption spectra that are much alike.
The problem is that on the basis of the absorption spectrum of an extract
of Sphaerobolus it would be difficult to identify a little riboflavin in the
presence of so much carotenoid pigment. An essential difficulty is that we
have no idea of the concentrations required for the system to work.
There is the further problem of a chemical receptor in relation to the
effect of orange light in the later stages of sporophore development. Some-
thing blue seems indicated. However, in extracts of sporophores at this
stage of sensitivity there appears to be no absorption in the visible region
above 520 nm. Here again the trouble may be connected with a very low
effective concentration.
In a culture of Sphaerobolus the orange colour is located in the sporo-
phores. When a dark-grown culture is transferred to light and sporophores
are initiated, the pigment increases and reaches a maximum about a week
before the first peak of discharge. It then falls off, but not to zero, to rise
again in anticipation of the next peak (Alloway & Ingold, 1971).
In addition to the orange hue of sporophores, cultures of Sphaerobolus
also develop a diffusible yellow pigment when grown on oatmeal agar.
Light is not necessary for its production. Its nature has not yet been
determined.
For sporophore production in Isolate A light and a temperature below
250 are vital external factors. There is, in addition, a third which, how-
ever, is unlikely ever to be limiting in nature. This is carbon dioxide. If the
air bathing a culture is deprived of its CO2 , no fruiting occurs. As with
light, the necessity for this factor obtains only during the first half of the
developmental process. Tracer experiments show that the C of the CO2 is
incorporated in the fungus (Ingold & Nawaz, 1967a).
Such a requirement is unusual in fungi, although it has been demon-
strated in Chaetomium (Buston, Moss & Tyrrell, 1966) and also in the
cultivated mushroom in which both vegetative growth and sporophore
initiation are dependent on carbon dioxide (Long & Jacobs 1969).
Turning to the question of the actual discharge of the glebal-mass, it is
well to have in mind the structure of the sporophore when fully grown but
before it has actually opened. Its histology has been the subject of detailed
study by a number of workers, especially Walker (1927) and Buller (1933).
The peridium has six recognizable layers (Fig. 9). Three are destined
to form the outer cup when the sporophore opens. Of the inner three
 Presidential address. C. T. Ingold 191

Fig. 9. Sphaerobolus stellatus. Vertical section of ripe but still un opened sporophore
(seated on its parent mycelial layer). An indicated sector is shown enlarged. The
peridium layers I, 2 and 3 will constitute the outer cup of the opened sporophore, and
layers 4 and 5 the inner cup. Layer 6 will undergo autolysis to produce the lubricating
fluid. G is the gleba, Very slightly modified after figures by Buller (1933).

layers, two go to form the inner cup, but the sixth, th e innermost, under-
goes autolysis to form the lubricating liquid in which, in the opened sporo-
phore, the glebal-mass lies loose but submerged.
The tissue active in discharge is the fifth layer. It is composed mainly of
elongated palisade cells stuffed with glycogen. However, in its upper part
this layer is pseudoparenchymatous and largely without glycogen. It is,
however, coloured orange with carotenoids which are also present in the
neighbouring part of the outer peridium (Alloway & Ingold, 1971).
The characteristic stellate opening of the spherical sporophore occupies
 Transactions British Mycological Society

South African t
isolate

A
t
~

IIsolate A
~mm I
I
I
I
I

Fig. 10. Sphaerobolus. The South African isolate and Isolate A, both to the same
scale, are shown at the moment of discharge, indicating for the former the inflow of air
as discharge occurs. To simplify the figure, the back half of the sporophore has been
omitted.

an hour or two, and once wide open, it is a matter of 5 or 6 h before dis-

charge occurs. During this time the glycogen in the motor region of the
inner peridial cup is converted to sugar, apparently glucose (Engel &
Schneider, 1963), to give a high osmotic pressure resulting in extreme
turgidity in the palisade cells. If this condition were to exist over the whole
area of the inner peridium, and particularly if it were to extend into the
orange teeth which attach it to the outer peridium, the whole membrane
would tend to tear away from its attachment on eversion. If this were to
happen regularly the discharge of the spore projectile might be much less
efficient.
Discharge is due to the sudden eversion ofthe inner cup. This has usually
been attributed to the sudden relief of tissue tension between the highly
turgid radial palisade cells tending to expand and the fibrous layer of fine
tangential interwoven hyphae forming the outer surface of the inner cup.
Buller (1933) has, however, suggested that only the inner layer is signi-
ficant and that the cup would still evert explosively even if the layer of
tangential hyphase were absent.
It is a significant fact that the inner cup is attached to the outer only
at certain points, so that before discharge the air between the two cups is in
free communication with the outside air. The final eversion of the inner
cup occupies only about a millisecond (Buller, 1933), so clearly there must
 Presidential address. C. T. Ingold 193
6 Isolate A

Of-- - - --t-----'----+-- - ..I.o---t-----''-----t

o 100 200 300 400cm
8 ,...-- ----, S. African isolate

c
z ') I------i

01--- --'----+-- - ...1..-- - 1--- - - - .,.-- - - - --,

o 100 200 300
Text-fig. II. Distance of glebal-massdischarge in Sphaerobolus. Bottom right: Spore-
mass mortar (tilted at 45° to vertical) used in studying distance of horizontal throw.
It consists ofa beaker lined with a roll offilter paper wetted by water in the beaker. The
'ammunition' is a culture in a 5 em diameter Petri-dish attached inside the beaker by
a blob of plasticine. Top: S. stellatus, isolate A: record of glebal-masses caught on each
50 em-long portion of a horizontal strip of white wallpaper at increasing distances from
the mortar. Bottom: The same but using the South African isolate. Glebal-masses
of the two isolates are shown to the same scale.

be a sudden inrush of air. The mechanism could hardly work if the two
cups were united completely by their rims (Fig. IO).
The maximum horizontal throw recorded for the glebal-mass of
Sphaerobolus is 569 em, a clear record for fungal gunnery with the sporan-
gium of Pilobolus longipes a poor second at 263 em and the ascospore-mass
of Podospora curvicolla a long way behind at 80 em (Buller, 1933). Individual
isolates of Sphaerobolus vary in the vigour of discharge. Thus for Isolate A a
maximum horizontal throw of 382 cm has been measured with a mean
value of 237 em. However, in the very large isolate from South Africa the
maximum was only 163 em and the mean 88 cm (Fig. II).
In my general experimental interest, I have been successively con-
cerned with a number of fungi. I dallied with Daldinia, an intriguing and
well-ordered fungus, but limiting since stromata have to be gathered from
dead ash trees. Then followed a Sordaria phase. Much was learnt from S.
fimicola, a reliable species easy to handle in the laboratory but nevertheless
rather inclined to mutate. During the past few years my work has been
largely concentrated on Sphaerobolus which is a joy to handle. It behaves
194 Transactions British Mycological Society
with clock-like regularity. Ifproperly treated, its capacity to fruit remains
undiminished and experiments are repeatable, even after a considerable
lapse of time, in a most satisfactory manner. I would commend it as re-
search material especially as it can be used in the study of so many aspects
of fungal physiology.
My period of work in a sophisticated laboratory is now very near its
end. I shall, indeed, miss my daily contacts with experimental work and
with those who have joined me in studies of spore discharge.
So ends my story of Sphaerobolus. So is my swan-song sung (Fig. 12).

c~~_
::- -==-
~ --=,,::=.:-:_:o-_~_,,,_,,,,_,,,_-_=-:,,"_,,.._:;:_:-_=- _-=. -=-_ _-_---.:_=__

Fig. 12

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ofBotany 27, 123- 145.
ALLOWAY, ]. & INGOLD, C. T. (1971). Physiological observations on Sphaerobolus,
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BULLER, A. H. R. (1933). Researches onfungi, vol. 5. London: Longmans, Green and Co.
BUSTON, H. W., Moss, M. O. & TYRRELL, D. (1966). The influence of carbon dioxide
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 Trans. Br. mycol. Soc. VoL 58. Plate 2I

r- - - - - -
Carpobolus Ja b .1'11

Frr;·
1
L

(Facing ,b. 194)

Trans. Br. mycol. Soc. Vol. 58. Plate 22

.., .
:.;
..

c
....
,
.....
"
' .

. .......,
"
'
"

'. '

.
"

"

"

Il,m
If
 Presidential address. C. T. Ingold 195
ENGEL, H. & SCHNEIDER, J. C. (1963). Die Umwandlung von Glykogen in Zucker in
den Fruchtkorpern von Sphaerobolus stellatus (Thode) Pers. vor ihrem Abschusz.
Berichte der Deutschen botanischen Gesellschaft 75, 397-400.
FRIEDERICHSEN, I. & ENGEL, H. (1957). Beitrage zur Kenntnis des Abschussrhythmus
und des Farbstoffs von Sphaerobolus stellatus (Thode) Pers. Planta 49, 578-587.
FRIEDERICHSEN, 1. & ENGEL, H. (1960). Der Abschussrhythmus der Fruchtkorper von
Sphaerobolus stellatus (Thode) Pers, Planta 55, 313-326.
HONDA, Y., SAKAMOTO, M. & aDA, Y. (1968). Blue and near ultra violet reversible
photoreaction on the sporulation of Helminthosporium oryzae. Plant andCellPhysiology 9,
603--60 7.
INGOLD, C. T. (1969). Effect of blue and yellow light during the later developmental
stages of Sphaerobolus, American Journal of Botany 56, 759-766.
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logical Society 56, 105-113.
INGOLD, C. T. & NAWAZ, M. (1967a). Carbon dioxide and fruiting in Sphaerobolus
Annals of Botany 3 1, 35 1-357.
INGOLD, C. T. & NAWAZ, M. (1967b). Sporophore development in Sphaerobolus: effect
of blue and red light. Annals of Botany 31,469-477
INGOLD, C. T. & NAWAZ, M. (1967c). Sporophore production in Sphaerobolus with
special reference to periodicity. Annals of Botany 31, 791-802.
INGOLD, C. T. & PEACH, J. (1970). Further observations on fruiting in Sphaerobolus in
relation to light. Transactions of the British Mycological Society 56, 105-113.
LONG, P. E. & JACOBS, L. (1969). Some observations on CO 2 and sporophore initiation
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MANACHERE, G. (1970). Recherches physiologiques sur la fructification de Coprinus
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phores. Annales des Sciences Naturelles, Botanique, Paris. 12" Serie, II, 1-96.
MICHELI, P. A. (1729). Nova Plantarum Genera. Florence.
NAWAZ, M. (1966). Sphaerobolus stellatus and light. Ph.D. thesis, University of London.
NAWAZ, M. (1967). Phototropism in Sphaerobolus. Biologia 13, 5-14.
RAMSBOTTOM, J. (1941). Dry rot in ships. Essex Naturalist 25,231-267.
SCHNEIDER, J. C. (1964). Der Farbstoff der Fruchtkorper von Sphaerobolus stellatus
(Thode) Pers. Planta 63, 351-355.
WALKER, L. B. (1927). Development and mechanism of discharge in Sphaerobolus
iowensis n.sp, and S. stellatus Tode. Journal of the Elisha Mitchell Scientific Society 42,
151- 178.

EXPLANATION OF PLATES
PLATE 21
Micheli's figure (1729) of his Carpobolus, a name changed by Tode in 1790 to Sphaerobolus.

PLATE 22
Sphaerobolus stellatus (Isolate A). (A) Gemma in L.S. with single germ-tube; from glebal-mass
12 h after its discharge. Note cellwall ofgemma is 0'08 /Lm thick, and that of the germ-tube about
half that thickness: L, osmiophilicgranules just inside the wall. The three large black bodies in
the gemma are contracted osmophilicbodies. (B) Detailsof the clamp separating the gemma from
its germ-tube: P, parenthosomes capping the dolipore. (C) Basidiospore from freshlydischarged
glebal-mass in L.S. Note cell wall (0'5 /Lm thick): N, nucleus; PM, plasma membrane.
Electronmicrographs by Dr Judith Alloway.

13 MYC 58