PLANT SCIENCE BULLETIN
A Publication of the Botanical Society of America, Inc.
1970 Volume Sixteen Number Three
Flowering and Photoperiodism of Plants1
the Fiftieth Anniversary of the Discovery of Phoroperiodism )
K. A. Timiryazev Institute of Plant Physiology of the Academy of Sciences
of the USSR (Moscow)
is a turning point, a cardinal period in the onto-genesis of all seed plants
when they turn from vegetative growth to generative development. The discovery
of photoperiodism, i.e., the response of plants to the duration of daylight
or to the length of day, was of particular importance in the sense of flowering
ecology and physiology (Garner and Allard, 1920; 1923). Day length appeared
to affect various aspects of ontogeny: growth, flowering, fruiting; the development
of above-ground organs—stems, and Ieaves, underground organs—roots,
tubers, bulbs, rhizomes; sexual differentiation, anatomical-morphological
structure, and the physiological processes related to all of these changes
(Samygin, 1946; Konstantinov, Sokolova, and Sakova, 1965).
influence of the length of day on the transition of plants from vegetative
growth to generative development is, however, the most significant and fundamental
phenomenon. Therefore, flowering was assumed by Garner and Allard to be the
main expression or criterion of photoperiodism. By the capability of plants
to flower under one or another length of day they are classified into various
photoperiodic groups; long-day, short-day, day-neutral, and others. Photoperiodism
appeared to be such a powerful and reliable, means of controlling the course
of plant development that after its discovery the entire subsequent investigation
of internal factors of flowering and of plant photoperiodism proceeded to
be inseparably linked.
this respect general phenomena observed during the early period of studying
the ecology and physiology of photoperiodism such as: the substantiation of
photoperiodism as an adaptive response, the differentiation of
1 This paper is a revised version of the Invited Paper presented
by Professor Chailakhyan at the XI International Botanical Congress, Seattle,
Washington, on August 28, 1969.
leaf and stem phases of the photoperiodic process, the clarification of the
importance of light and darkness, and the demonstration of trophic and hormonal
factors which form the basis of photoperiodism, are of great importance (Chailakhyan,
1967; 1968). In the same connection the following more recent theories and ideas
on plant flowering, that are at the same time theories and ideas on photoperiodism,
are gaining great interest: 1) the theory of endogenous rhythms (Bunning, 1950.
1956, 1958), the basis of which is formed by the idea of the relation of photoperiodic
response to the diurnal endogenous rhythm; 2) the phytochrome theory (Borthwick,
Hendricks, Parker, 1952), on the dependence of the course of photo-periodic
response on a special pigment, phytochrome, that has two mutually interconvertible
forms; 3) the hypothesis on the correlation of the rates of light and dark responses
(T.yubimenko and Scheglova, 1927; Chailakhyan, 1956; Konstantinova, 1966) which
is distinct for long-day and short-day species; 4) the conception of the two-phase
character of flowering: a) the phase of flower-bearing stem initiation and b)
the phase of flower initiation (Chailakhyan, 1964).
is yet no solution to the basic mystery of photoperiodism: why do typical
long-day plants flower under a long day and fail to do so under a short day,
while short-day plants flower under a short day and fail to do so under a
long day? The diametrically opposite response of long- and short-day species
to length of day, the duality in their behavior, as Maksimov (1925) wrote
in his review on photoperiodism, has surprised everyone and still remains
mystery is deepened by the fact that it was demonstrated that physiological
processes and biochemical reactions which precede flowering respond to length
of day in the same manner in long- and short-day species. The increase in
activity of metal-containing enzymes and of the content of carbohydrates and
growth hormones in all plants regardless of the nature of their photoperiodic
response proceeds more intensively under long days, while the increase in
activity of residual respiration enzymes and in content of nitrogenous compounds,
the products of nucleic metabolism, and substances of the anthesine type that
stimulate floral initiation proceeds more intensively under short days (Chailakhyan,
carried out recently in the Laboratory of Growth and Development of the Institute
of Plant Physiology of the Academy of Sciences of the USSR have shown that
light, darkness, and length of day affect
PLANT SCIENCE BULLETIN
ADOLPH HECHT, Editor
Department of Botany
Washington State University
Pullman, Washington 99163
Harlan P. Banks, Cornell University
Sydney S. Greenfield, Rutgers University
Robert W. Long, University of South Florida
William L. Stern, University of Maryland
Erich Steiner, University of Michigan
December 1970 Volume 16
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activity of oxidation-reduction processes and the coupling of respiration
and phosphorylation in the same direction in the long-day as well as in the
photochemical activity of chloroplasts was always greater in all plants under
long-day conditions regardless of their photoperiodic response (Bavrina, 1969).
The coupling of respiration and phosphorylation in the plants of both mentioned
groups changes in the same manner according to the Iength of day: during the
daylight hours the coupling increases; under darkness, particularly during
a lengthy night of the short-day cycle, the coupling gradually decreases (
Aksenova, Konstantinova, Nikitina, 1968) . It appeared that uncoupling of
respiration and phosphorylation during a short-day cycle, achieved by introducing
2,4-dinitrophenol into plants, failed to affect flowering under light conditions
while under darkness changes the course of dark responses; the inhibitor fails
to produce marked changes in the long-day cycle whether applied during the
light or dark period. Application of diuron, an inhibitor of photophosphorylation,
has led to substantial changes in the flowering of plants under long days,
where light responses play a determinative role, but flowering was unaffected
under short-day conditions where darkness responses are of primary importance
(Konstantinova, Aksenova, Nikitina, 1968; Bavrina, Aksenova, Konstantinova,
1969). Thus, light responses of the photo-periodic process in plants of both
photoperiodic groups require energy of photophosphorylation while dark responses
require energy of oxidative phosphorylation (Chailakhyan, Aksenova, Bavrina,
a result, changes in physiological and biochemical processes and in nutrient
and regulatory substances in all plants occur under the influence of the length
of day in the same direction. On the other hand, some plants flower under
a long day and others under a short day. Concluding that the conversion to
floral initiation in long- and short-day species is preceded by opposite metabolism
is entirely unsuitable from the general biological point of view. Yet, one
has to admit that the investigation of physiological and biochemical processes
affected by the length of day has not decreased the mystery of photoperiodism.
there are approaches to the solution of this mystery: an historical or evolutionary
approach that offers an ecological explanation of the formation of various
photoperiodic groups including long- and short-day species, and an experimental
approach which draws us nearer to the physiological explanation of the opposite
behavior of these species.
The Genesis of Photoperiodic Groups
historical or evolutionary approach is based on in. vestigations of photoperiodic
response of various plant species and cultivars distinguished by their origin
and ecology. Experiments showing close relation between the character of photoperiodic
response and the geographical origin of plants have shown photoperiodism to
be an adaptive response. Moreover, specialization of plants growing in temperate
and high latitudes tends toward the formation of long-day species incapable
of flowering under a short day, and the specialization of plants of subtropics
and tropics is directed towards the establishment of short-day species (Lyubimenko
and Scheglova, 1927; Doreshenko and Razumov, 1929; Allard, 1932; Konstantinov,
1934; Razumov, 1954). Short- and long-day species originated from day-neutral
species which are more ancient by origin, and the adaptation of the former
species to length of day gave them advantages over neutral forms. The fact
that long-day plants fail to flower under a short autumn and winter day but
instead produce small bushes and rosettes promotes their preservation under
a snow cover and represents their adaptation to overwintering in temperate
and northern latitudes. Short-day plants fail to flower under long summer
days but continue vegetatively and survive hot and dry summers or periods
of pouring rains in subtropical and tropical countries.
adaptive significance of photoperiodism goes far beyond controlling the dates
of flowering. It plays an out-standing role in permitting individual or species
survival under adverse environmental conditions. The changing length of day
gives the plants signs of the necessity of preparing for seasonal changes.
The length of day, as an astronomical phenomenon, is a reliable and accurate
guide pointing out to plants when they should flower and re-produce- and when
to be prepared for adverse environmental conditions (Katunsky, 1939; Moshkov,
1940; Samygin, 1946; Biinning, 1948; Chailakhyan, 1956; Skripchinsky, 1958,
conversion to generative development does not depend only upon the variation
of the length of day. The flowering in a large group of species neutral to
the length of day and failing to exhibit photoperiodic adaptation is governed
by other environmental conditions—temperature, intensity and quality
of light, nutrition and water regime. The response of these species to environ-
conditions is also adaptive and permits the con-version of neutral plants
to the stage of sexual maturity. The same environmental conditions also play
an important role in the flowering process of photoperiodically sensitive
species, because the photoperiodic response of long-day, short-day, and other
species may occur only under a definite level of temperature, light intensity
and quality, nutrition and water regime.
Long,-short-day species (Bryophyllum, Kalancboe, A1oë,
Cestrum, etc.) are known. They flower only under successive exposure
to long and then to short days (Dostal. 1949, 1950; Resende, 1952, 1954; Sachs,
1956). There are also short; long-day species (Scabiosa, Campanula, Tri f oliur,
etc.) which flower only under successive re-placement of short days by long
days (Chouard, 1957; Wellensiek. 1960: Thomas, 1961 ). Medium-day (stenophotoperiodic)
and extreme day length. (amphiphotoperiodic) species are also known. The former
flower under an intermediate day and the latter flower only under a long day
or a short day but fail to flower under an inter-mediate day (Lang, 1965). The
adaptive character of the plant response to the length of day which is more
general in long- and short-day species and more specialized in other species
is especially distinctive under this classification.
photoperiodic response of plants was initially considered as a static response
to permanent short or long day lengths. The classification into long- and
short-day species was related to the same concept. But it soon turned out
that the alternating length of day which occurs in nature is of great importance.
Thus, McClelland (1924) showed that Tephro.tia candida, a tropical species,
flowered only under a 12-hour day. Flowering is even and abundant if the plants
previously were exposed to a 13.5-hour day and, if the plants were first exposed
to a 10-hour day, flowering proceeds poorly or not at all.
adaptive response to day length is very pronounced in long,-short-day and
short,-long-day species. In the former this response means an adaptation to
the changing length of day in the summer-fall season. In the latter it means
an adaptation to the changing length of day in the spring-summer season. The
presence of ecotypes with various photoperiodic responses within the same
population should also be regarded as an adaptation to changing daylength
of the growing season (Sinskaya, 1960). All this indicates the dynamic character
of photoperiodic adaptation.
adaptation has developed along with the formation and establishment of various
plant forms with morphological features and physiological characters peculiar
to them. Photoperiodic adaptation is representative not only of the angiosperms
but of lower plants, including fungi and algae. There have been developed
species of angiosperms with a quantitative photoperiodic response, i.e., capable
of flowering under any length of day, but with some acceleration under long
days in long-day species and with some acceleration under a short day in short-day
species. Species with a qualitative photoperiodic response, typical long-day
species capable of flowering only under long days, and typical short-day species
capable of flowering only under short days that have lost their responsiveness
under usual conditions have been developed.
question of what morphological and physiological changes are related to the
loss of flowering ability of long- and short-day species is of substantial
importance in the approaches to the solution of the main mystery of phoroperiodism.
Adaptation of typical long-day species to overwintering is related to their
loss of ability for stem formation and growth, while the adaptation of typical
short-day species to existing through the summer period of tropical rains
and lasting drouths is related to their loss of the ability to produce flowers
(Chailakhyan, 1960, 1961). The separation of the process of flowering into
two phases occurs in both cases: a phase of flower-bearing stem formation
and a phase of floral initiation. Formation of flower-bearing stems in long-day
species proceeds more rapidly or only under a long day while the floral initiation
in short-day species proceeds more rapidly and only under a short day.
the loss of ability to flower under an adverse length of day in photoperiodically
sensitive species occurs not completely but partially: in long-day species
it is the loss of the ability of stem formation and growth under the short-day
conditions, in short-day species it is the loss of the ability of floral initiation
under a long day.
Conception of the Two-Phase Character of Flowering
comparative experimental investigations on the role of trophic and hormonal
factors in the flowering of long- and short-day species and the results of
the experiments on the genesis of photoperiodic groups enabled us to set up
an idea about the two-phase character of flowering (Chailakhyan, 1964). Each
of these morphological phases corresponds to a certain physiological condition.
The first phase is characterized by intensive production of gibberellins in
leaves accompanied by an intensification of carbohydrate metabolism and the
rate of respiration at the expense of metal-containing enzymes and by an increase
in auxins in stem buds. The second phase is characterized by intensive production
of anthesins in leaves, which are essential for floral initiation. This process
is accompanied by the intensification of nitrogenous metabolism and of the
activity of residual respiration as well as by an increase in nucleic acids
(see Table 1_) .
content of gibberellins plays a decisive role for physiological changes in
the first phase of flowering. Anthesins that affect the floral initiation
(not yet isolated) play a decisive role in the second phase of flowering.
The two phases in day-neutral species proceed regard-less of the length of
critical phase in long-day species is the first one. the phase of flower-bearing
stem formation, which proceeds only under a long day ("L") when the intensive
production of gibberellins occurs. Capacity for the second phase of flowering
in long-day species is firmly fixed just as in day-neutral species, and it
proceeds equally under a long as well as under a short day. The second phase
of flowering in long-day species is nor critical because they initiate flowers
under a short day after a gibberellin treatment of central buds.
second phase of flowering, flower initiation, is critical in short-day species,
and it proceeds only under a short day. The ability to conduct the first phase
of flowering in short-day species is firmly fixed just as in day-neutral species
and proceeds equally under a long as well as under a short day. The first
phase of flowering in plants of short-day species is not critical because
these species produce sterns under long as well as under short days.
this suggests that although the course of physiological and biochemical processes
proceeds in both photo-periodic groups in response to the length of day in
the same direction, the results are different. Metabolic changes occurring
in response to long days that determine the events of the first phase of flowering,
stem formation and growth, are important for the flowering of long-day species;
the occurrence of the second phase, flower initiation, is ensured by metabolism
which is controlled not by the length of the day but by immediately translated
hereditary information. The metabolic changes in response to a short day determine
the second phase of flowering and are important for the flowering of short-day
species; the passage of the 'first phase, stem formation and growth, is ensured
by metabolism which is controlled not by the length of day but by the immediately
read genetical information. Beyond this control there are various phases of
flowering and biosynthesis of various constituents of the two-components system.
of the flowering hormones in the two photoperiodic groups (Chailakhyan, 1967,
1968; Chailakhyan, Aksenova, Bavrina, Konstantinova, 1969).
Factors and Internal Factors
discrepancy and duality of photoperiodism of long-and short-day species are
completely determined by the discrepancy and duality of the very criterion
of photoperiodism, i.e., flowering. This criterion is distinct in reference
to the two groups of long- and short-day species, the difference of the criteria
being determined not only by environmental factors, that is factors by the
length of the day, but also by internal factors.
neutral species have no adaptation to the length of day, and their passage
through both phases of flowering is related to direct translation of genetic
in-formation. If the neutral response in reference to the length of day is
marked by the letter "N," the long-day response by the letter "L," and the
short-day response by the letter "S," then the symbol for neutral species
can be assumed as "NN."
species, by virtue of adaptation to environmental conditions, are long-day
ones only in the first phase; in the second phase they are neutral ("LN").
Short-day species remain photoperiodically independent of length of day for
the first phase; in the second phase, coming under the control of the environment,
they are short-day ("NS"). Long,-short-day and short,-long-day species have
followed the course of adaptation to the length of day still further: both
phases of flowering are under the control of environment and are determined
by the length of day. Consequently, long,-short-day species can be marked
by letters "LS" and the short,-long-day species can be marked by the letters
"SL" (Chailakhyan, 1969). The first phase of flowering in long,-short-day
species proceeds only under a long day while in short,-long-day species this
phase proceeds only under a short day. This is puzzling because the first
phase of flowering is associated with the controlling action of a long day
(in long-day species) or with control independent of the length of day (in
of changes caused by age and of the juvenile stage of long,-short-day species
enables us to draw certain conclusions about the two-phase photoperiodic control
of the flowering of plants. It is known that the period of formation of the
vegetative organs before the onset of flowering, or the period of the juvenile
phase, fluctuates between a few days and a few months (Lang, 1965). The flowering
response of long,-short-day Brynphyllum, which sets in at a successive replacing
of long days by short days, occurs only in adult plants at the age of no less
than one year with no less than 10-12 pairs of large well-developed leaves
(Resende, 1952; Zeevart, 1962). Younger plants fail to flower under such a
manipulation of the photoperiodic regime.
is possible to reduce this long period of the juvenile phase by treating the
plants with gibberellins. It was demonstrated that Bryophylium plants thus
treated flower under constant short days (Biinsow and Harder, 1956; .Biinsow,
Penner, and Harder, 1958). Subsequently it appeared that three-month-old plants
flower as a result of treatment with gibberellin (Resende and Viana, 1959;
Penner, 1960; Wadhi and Mohan Ram, 1967) .
our experiments plantlets formed from adventitious foliar buds of Bryophyllunt
daigremontianum initiated flower buds 50 days after spraying with gibberellin
under short days. Young shoots formed from such buds that were still undetached
from rooted leaves flowered under the same conditions (Chailakhyan, Yanina,
Frolova, 1968, 1969).
the period of the juvenile phase is shortened by gibberellin and the process
of flowering sets in under the influence of short days, then according to
Zeevart (Zeevart, 1969), Bryophyllum can be considered as a typical short-day
species ("NS"). However, in contrast to short-day species in Bryophyllum and
apparently in other long,-short-day species, the juvenile phase is con-trolled
by the length of day and proceeds only under long days.
data are absent for short,-long-day species (Scabiosa, Campanula, and Coreopsis)
, but it is known that these species under short days are in the rosette or
bush form and start to produce stems only when exposed to long days (Chouard,
1957; Wellensiek, 1960; Ketellapper and Barbaro, 1966). They may thus be considered
typical long-day species ("LN"). But the juvenile phase in short,-long-day
species in contrast to long-day species is controlled by the length of day
and proceeds only under short days.
juvenile phase in short- and long-day species of annual plants is independent
of the length of day in spite of the fact that in certain forms, as in tobacco,
the short-day Mammoth (N. tabacunt) and the long-day Sylvestris (N. sylvestris),
the juvenile phase is of long duration (Yevtushenko, 1947). The photoperiodic
response of long,-short-day and short,-long-day species is distinguished by
the fact that their juvenile phase is also under the control of the environment.
Long,-short-day species actually are short-day ones: in the first phase of
flowering they are neutral and in the second phase they are short-day ones
but their juvenile phase proceeds only under long days ("L-NS"). Long,-short-day
species basically are long-day ones: in the first phase they are long-day
species; in the second phase they are neutral and their juvenile phase proceeds
only under short day ("S-IN").
adaptive significance of controlling the juvenile phase by exposure to long
days consists in the prevention of too-early flowering of young plants (Chailakhyan
and Podolny, 1968). This phenomenon in long,-short-day species is seen in
the fact that leafy plants with large stems are formed under long days prior
to the conversion to flowering while in short,-long-day species it is seen
that, prior to the conversion to flowering, vigorously tillering forms with
numerous leaves and stems develop under short days.
data on the relationship of the juvenile phase and the flowering phases, both
controlled by the conditions of the length of day and independent of the latter
in the plants of various photoperiodic groups, are summarized in Table 2.
the flowering of plants of various photoperiodic groups falls under the controlling
action of the length of day to a variable degree: in long- and short-day species
a lesser extent while in long,-short-day and short,-longday species to a greater
extent. The extent is determined by the adaptive function of photoperiodic
response which permits the plant to flower at the most favorable age and season
Control of the duration of the juvenile phase in plants occurs not only under
the influence of the length of day but also of other environmental and internal
factors. Here it is essential to point out that the passage of the juvenile
phase in some short,-long-day species can be effected by exposing them to short
days or by treatment with lower temperature (Lang, 1965). The juvenile phase
of winter forms and biennials is controlled by lower temperature, and since
they behave as typical long-day species in subsequent responses (Chailakhyan,
1942), they can be denoted as "T-LN." The juvenile phase of perennials, characterized
by a significantly longer duration than in annuals, is controlled by changes
caused by age and internal factors.
The Mystery of Plant Photoperiodism and the Outlook of Its Solution
of environmental factors and of the possible role of internal factors in the
conversion of plants to flowering enables us to approach the solution of the
mystery of photoperiodism in long- and short-day species.
as a main criterion of photoperiodism is entirely different for these two
types. In long-day species flowering is reflected by the formation of flower-bearing
stems under long days and is related to the production of gibberellins and
other changes in metabolism. In short-day species flowering is reflected by
flower initiation which proceeds under short days and is related to the production
of substances of the anthesin type and other changes in metabolism (Table
I ). The production of gibberellins and anthesins and the course of other
biochemical processes related to the length of day are entirely the same in
both types. The resulting effect, however, is different because in long-day
species the genetically fixed ability to produce gibberellins was lost, and
balanced by the physiological process of their production under long days.
The ability of producing substances of the anthesin type was lost in short-day
species, and this loss is balanced by the physiological process of their production
under short days. This phenomenon was demonstrated for long-day species (Lang,
1956, 1957; Marth, Audia, and Mitchell, 1956; Biinsow and Harder, 1956, 1957;
Lona, 1956, .1957; Chouard, 1957, 1958; Chailakhyan, 1957, 1958, and others),
for Iong,-short-day species (Biinsow, Penner, and Harder, 1958; Resende and
Viana, 1959; Penner, 1960) as well as for short,-long-day speices (Chouard
1957; Ketellapper and Barbaro, 1966) by the insertion of gibberellins under
the conditions of short days. Probably the loss of genetically fixed ability
for intensive production of substances of the anthesin type in short-day (NS),
long,-short-day (L-NS), and short,-long-day (S-LN) species under any length
of day could also be substituted by the insertion of sub-stances of the anthesin
type under the conditions of long days. However, such substances have not
been isolated from plants, and conclusions about them are drawn on the basis
of indirect biological responses.
genetic information about the process of flowering in species that are neutral
to the length of day is realized, regardless of, the length of day, directly
through an internal hereditary program. The hereditary control of flowering
which is independent of the length of day can be called autonomous control.
genetic information about the process of flowering in species adapted to the
length of day is also translated through an internal hereditary program while
its direct realization is put under the control of an environmental factor,
the length of day. This genetic control of flowering that was formed in the
process of evolution can be called photoperiodic control (Table 3).
as seen in Table 3, the properties controlled by daylength and those properties
not under such control interact in various combinations with the photoperiodic
phenomena and with the genetic complement of the plant. These properties do,
however, change. If the production of floral_ hormones by day-neutral plants
is under autonomous control, then as adaptation to daylength progresses, hormone
production will increasingly come under photo-periodic control in the evolving
long- and short-day plants. Ultimately, hormone production will be completely
under photoperiodic control in long-short-day and short,-longday species.
recognition of the role of factors related to both photoperiodic and autonomous
control permits us to come closer to the solution of the main mystery of photoperiodism
and to the analysis of the phenomenon as a whole. Two trends in further investigations
appear. First, the identification and description of the mechanism of action
of hormonal factors in various phases of conversion of plants from vegetative
growth to flowering in connection with photoperiodism and other adaptive responses
as well as changes in plants caused by their age. Secondly, the identification
and description of the mechanism of the realization of genetic information
that determines the flowering of plants by using the outstanding progress
achieved in studying the heredity of plants (Bonner, 1968) and in connection
with investigations on hybridization of long- and short-day species as well
as other ecotypes that sharply differ in the dates of flowering (Herer, 1950;
Naylor, 1953; Bremer, 1961; Griesel, 1966), on polyploidy (Konstantinov and
Zhebrak, 1963) ; and on plant mutagenesis.
in these two directions bound by a common and single goal seem to be the most
promising and would be able to initiate a new semicentennial history in the
study of photoperiodism.
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Ceskoslov. Akad. Zemedel. 22, 241; 1950.
Acad. Sci. Nat. Moravosiles (Brno), 22, 3, 57.
W. W., Allard, H. A. 1920. J. Agric. Res., 18, 553;
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A. 1966. (Dyton, 23, 33.
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N. N., and Zhebrak, E. A. 1963. Dokl. Akad. Nauk SSSR, 150, 5, 1149.
N. N., Sokolova, T. N., and Sakova, T. V. 1965.
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61, 118. Konstantinova, T. N., Aksenova, N. P., and Nikitina, A. A.
Fisiol. Rast., 15, 4, 631.
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Sci. U.S.A., 43, 709; 1965. Encyclopedia of Plant Phy-
15(1): 1380 (Ruhland, W., Ed., Springer, Berlin). Lona, F. 1956. L'Anteo Parmense,
27, 867; 1957. 28, 111. Lybimenko, V. N., and Scheglova, O. A. 1927. The Journal
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P. C., Adia, W. V., Mitchell, G. W. 1956. Bot. Gaz. 118, 106.
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M. 1959. Porfug. acta biol. YI, 77. Sachs, R. M. 1956. Plant Physiol, 31,
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R. G. 1961. Nature, 190, 1130.
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J.A.D. 1962. Planta 5S, 543; 1969. 84, 339.
Botany Is Not Dead—It's Still Sleeping1
Office of Biological Education
Hark! The jolly green giant mutters in his sleep. When William Stern's Quo
Vadis Botanicum? crossed my desk
The title and the paragraphs that follow it are offered as one man's opinion
and should not be construed as representing the opinions of his employer.
knew it would be followed by a series of remarks like those in the December
issue of Plant Science Bulletin. Having been a botanist all my professional
life, I was all too familiar with that type of wailing. It's almost diagnostic
of a botanist that he blames his second-class citizen-ship on almost everyone
but himself. Blame the trend toward integrated biology departments. Blame
CUEBS for its concept of a core curriculum for all of the biological sciences.
Blame the AIBS for succeeding at bringing some order into the organizational
chaos of the American Biological Community. When all else fails, blame the
unenlightened administration. There are plenty of targets. Have at it. Play
Don Quixote while the giant sleeps on.
real reason for our dilemma was much more accurately described by jack Sharp
during a recent visit to AIBS. The plain fact is that botanists have failed
miserably at convincing the public or even the hordes of students they process
that plants are important, that plants hold the sine qua non of survival,
and that botany is where much of the action must be if man is to share in
this survival. Yes, the plain fact is that botanists, you and I, have failed.
It's not really surprising, however, when you see the way we have strutted
around uttering statements about the importance of botany which were supposed
to be true simply because we said them. Plants are important, there is no
argument on that. It remains to be seen whether botanists will be.
Galston's otherwise fine article (Plant Science Bulletin 16(1) :1-7) seems
to miss the main point when he says,
I think the first thing we need to tell our young friends who are challenging
us to climb down out of the ivory tower into the cobblestone strewn streets
is 'Fine, we will send you a delegation; but some will have to stay up here
to keep the store running.'
delegation (or is it relegation) should be to the tower not the streets. If
we botanists don't get our people into the streets to learn how to get their
hands on the levers of social change and political responsiveness we will
see continuing reduction of support for those few creative hothouse flowers
who should remain in the tower, if indeed by some cultural accident the tower
is left standing. Whether the omniscient "we" in the above quote are really
able to "tell our young friends" anything remains to be seen. We may be beyond
telling them anything; we may have to show them. If all we can show them is
a delegation, it had better be a darned large one. They (the students, the
people) have been put-off and put-down by the tower and its self-serving rhetoric
far too often. They aren't going to stand for much more of it. When Galston
indicates that he believes in "the collective wisdom of the people of a democracy
to decide important matters concerning their own fate" he gets at the real
problem. The "collective wisdom" is rapidly reducing budgets and increasing
its call for relevant performance with the little money that can be spared
from the higher priority jobs we have too long ignored while strutting around
in our towers. We had better come down from our. "ego trip" and get to work—real
work. Or is it all just a dream?
has the giant done to bring on his present ap-
stupor? For years he's cranked out papers in his pct esoteric area with one
hand and graduate students with the other while the real world was falling
apart at the seams. Scan the last few issues of most botanical journals and
see how much you find that is really relevant to the salvation of the planet
and the unfinished business of mankind. Sure, there will be a few papers scattered
here and there that have relevance to real problems but they will likely be
obscured by great masses of pap on The Research I Did Last Summer," The Cartwheels
My Students Will Turn In Order To Get Their Degrees," or worst of all, "I
Just Kept Turning The Same Old Crank And This Is What Came Out." Please don't
misunderstand, the problem isn't with the journals, it's with the heads of
the botanists who will feed and put up with this sort of stuff—they're
cheered when I heard Arthur Cronquist present a resolution before an AIBS
Governing Board Study group to the effect that the most important concept
we must impart to our students is that of spaceship Earth with its finite
resources, limited channels for the recycling of materials, and serious people
pollution problems. On the 26 campuses I've visited this year and last, I
have not seen too much of that going on. The green giant is still hung up
in the plurilocular zoosporangia of Ectocarpus, the intercalary meristems
of one grass or another, or trying to get the correct number of carbon atoms
in ATP while begrudgingly putting it on the blackboard. All of these things
and many more can be relevant. As dealt with, however, in most research and
teaching, they simply aren't.
time we realize that the research we do should be related to, if not aimed
at, real problems rather than our esoteric whims. The evidence is in. The
world at large and our students in particular have tuned us out. There is
an amazing amount of brain power in the botanical community. Every bit of
it is critically needed. Let the doubters look at the report by Steinhart
and Cherniak issued by the Executive Office of the President under the title
"The Universities and Environmental Education," or chew on the 1968 words
of the then U.S. Commissioner of Education, Harold Howe, who said, "Entirely
too many professors in graduate schools are interested only in the kind of
teaching which produces more professors in graduate schools."
it happens that there are botanists in graduate schools and elsewhere fighting
to prevent more Lake Erie messes; there are botanists researching perilously
important questions; and there are botanists telling it like it is to their
students. It's too bad they are so few in number that they constitute the
exception rather than the rule.
hark, the green giant is muttering! He is still asleep to be sure, but there
are signs of hope all around that one day soon he will wake up and take hold.
It isn't necessary to play at blaming everyone and everything else for his
long sleep. It is necessary to accept the blame ourselves and make a renewed
commitment to make our individual lives count in the battle for survival,
growth, and enlightenment. Botanists could become important yet!
NOTES FROM THE EDITOR
am pleased to announce that Dr. Robert W. Long, Professor of Botany and Chairman
of the Department of Botany and Bacteriology of the University of Southern
Florida, Tampa, has accepted appointment as the new Editor of the Plant Science
Bulletin. Dr. Long will begin his five-year term with Volume 17, the first
number of which will appear early in 1971.
Corrigenda: In the article "Terminology in the Plant Sciences" by W. C. Baum
(Plant Science Bulletin 16, No. 2), the term pantonenaatic should have appeared
at the end of the sentence on page 3, as follows: "3) flagella bearing lateral
appendages: pantonematic." Both J. L. Strother and W. C. Steere have called
attention to the fact that the term infructescence as used in the article was
NEWS AND NOTES
1971 Botanical Society Meeting
advance information is being provided so that you may begin making your plans
for the coming year. More detailed information is being prepared for our December
issue. In place of our usual late August meeting in association with other
AIBS societies, the Botanical Society of America will hold its annual meeting
in June, 1971 in collaboration with the Canadian Botanical Association. General
arrangements, however, will be handled by the AIBS. The following outline
has been provided by Dr. S. N. Postlethwait, our Program Chairman:
Our Northern Plants: Their Importance in the World's Resources
University of Alberta, Edmonton, Alberta Time: June 20-24, 1971
Chairman: Wilson Stewart
Alberta, Canada Probable deadlines:
of Field Trips and Symposia—December
of paper titles and abstracts—February
date for preregistration—May 15 Fees: Preregistration Late
Member $12.00 $15.00
Spouse 3.00 3.00
Student 5.00 8.00
Botanic Gardens at Bogor Damaged by a Hurricane
freak hurricane on January 4, 1970 resulted in extensive damage to one of
the worlds' leading botanical establishments. The loss to botany of such a
famous and important collection transcends national boundaries and
is felt that many of the institutions and individuals who have benefited from
the Gardens and its associated facilities such as the Treub Laboratory and
the Herbarium Bogoriense may wish to do something tangible to help.
University of Wyoming Offers Ph.D. in Botany
its regular August meeting, the Board of Trustees of the University of Wyoming
approved a Ph.D. program for the Department of Botany. Although the Rocky
Mountain Herbarium is recognized nationally and inter-nationally and a number
of well-known botanists have earned masters' degrees at Wyoming, the Ph.D.
has never been offered. Recently the faculty has been expanded to six full-time
members, and another appointment is anticipated for the 1971-72 academic year.
With the opportunity to pursue studies leading to the doctorate, students
with an interest in the area of environmental and evolutionary botany should
find Wyoming even more attractive. Recently remodeled laboratories and new
equipment pro-vide additional facilities for chemotaxonomic and biosystematic
research. For additional information, write: Dr. John R. Reeder, Head; Department
of Botany; University of Wyoming; Laramie, Wyoming 82070.
Major Evolutionary Events Symposium
page 6 of our June issue a preliminary announcement concerning publication
of the proceedings of this symposium appeared. The latest and more complete
information is the following:
Cambridge Philosophical Society is publishing, as Vol. 45, No. 3 of Biological
Reviews, the Symposium on "Major Evolutionary Events and the Geological Record
of Plants" presented at the XI International Botanical Congress, Seattle,
Washington, U.S.A., in September, 1969 under the Chairmanship of Professor
Harlan P. Banks. The Symposium is a summary of major events during a three-billion-year
span and will be of interest to a variety of readers. The separate papers
are as follows:
W. Schopf—Precambrian microorganisms and evolutionary events prior to
the origin of vascular plants.
G. Chaloner—The rise of the first land plants.
B. Beck—The appearance of gymnospermous structure.
Pettitt—Heterospory and the rise of the seed habit.
Muller—Palynological evidence on early differentiation of angiosperms.
P. Banks—Summary and time scale.
Symposium will be sold at the normal price of single issues of Biological
Reviews, :£2.12.6 (S8.00). This issue may be purchased through a bookseller
or through the American Branch of Cambridge University Press, 32 E. 57th Street,
New York 11022.
Otto Stein, Head of the Department of Botany at the University of Massachusetts,
announces the following
in his department: Albert C. Smith, formerly Wilder Professor at the University
of Hawaii, has been appointed as Ray Ethan Torrey Professor of Botany: Paul
J. Godfrey, formerly at Cape Lookout National Seashore, Beaumont, North Carolina,
has been appointed an Assist-ant Professor in the Department, as have Livija
Raudzens, formerly at Columbia University, and Peter L. Webster, formerly
at Brookhaven National Laboratories. John L. Harper, Head of the School of
Plant Biology of the University College of North Wales, and D. Ross Moir of
Branton University, Branton, Manitoba will be Visiting Professors. Professor
S. Shapiro will be on sabbatical leave to work at Wageningen in the Netherlands.
V. Raghavan, who has been at the University of Malaya, Kuala Lumpur, for the
past few years, will join the Academic Faculty of Botany, College of Biological
Sciences, Ohio State University, Columbus, as of the fall of 1970.
D. Voth became Professor Emeritus of Biology at the University of Chicago
in 1970. Dr. Voth will now be associated with the Department of Biological
Sciences, Northern Illinois University, DeKalb, Illinois.
K. Bonde of the University of Colorado has been named a visiting scientist
to Rumania under the U.S. National Academy of Sciences' Scientific Exchange
Pro-gram. Dr. Bonde is interested especially in Rumanian botanical studies
of Carpathian Mountain alpine regions, in relation to his Colorado Rocky Mountain
research on the growth and development of alpine plants.
William H. Matchett, formerly of the Battelle Seattle Research Center, has
been appointed Professor of Botany at Washington State University, and replaces
Adolph Hecht as Chairman of the Department of Botany. After 15 years as departmental
chairman, Hecht will continue as Professor of Botany and Genetics. Dr. Joseph
L. Hindman, formerly of the Department of Botany at the University of Georgia,
has been appointed Chairman of the Program in General Biology and Associate
Professor of Botany at Washington State. Dr. James C. Hickman, Assistant Professor
of Botany and General Biology, has resigned to accept a position at Swarthmore
Sterling B. Hendricks, research scientist with the U.S. Department of Agriculture
for more than 40 years, retired on July 3], 1970. Dr. Hendricks is best known
for his pioneering work in photoperiodism, and recently for his and his associates'
discovery and isolation of phytochrome, the protein molecule that regulates
many plant growth processes. Among the many awards that have been granted
in recognition of Dr. Hendricks' achievements are the Hillebrand Prize, the
Science Award of the Washington Academy of Sciences, the Day Medal, USDA's
Distinguished Service Award, the first Presidential Award for Distinguished
Civilian Service, the first Rockefeller Public Service Award, the Hoblitzelle
Award in Agricultural Sciences (with H. A. Borthwick), the Stephen Hales Award
(with H. A. Borthwick), the Distinguished Service Award of California Institute
of Technology, three honorary degrees, presidencies of several professional
societies, and membership in the National Academy of Sciences.
Neville Jones 1904-1970
Neville Jones, professor of botany and curator of the Herbarium at the University
of Illinois, died June 25 at the age of 66. He never recovered from injuries
received in an automobile accident on April 14, 1968, while enroute to Allerton
Park where he had collected plants for two decades.
numerous publications include the books, "Vascular Plants of Illinois," "Flora
of Illinois," "Annotated Bibliography of Mexican Ferns," and "Taxonomy of
the American Species of Linden."
January 26, 1904, in Boston, England, he moved to the United States with his
parents. He received his BS degree from the State College of Washington, Pullman,
in 1930, and his MS and Ph.D. from the University of Washington, Seattle,
'in 1932 and 1937. From 1937 to 1939 he was on the faculty at Harvard University
as instructor in biology, tutor in the division of biology, and technical
assistant in the Arnold Arboretum. He came to Illinois in 1939.
1967, Dr. Jones received the Eunice Rockwell Obberly Award from the American
Library Association for his "Annotated Bibliography of Mexican Ferns," judged
the best bibliography in the field of agriculture or related sciences. He
was a Guggenheim Fellow in 1944, a visiting lecturer at the University of
Wisconsin in 1947, and a visiting lecturer at the University of Colorado in
the summers of 1960 through 1965. From 1944 through 1961, he was editor and
reviewer for the American Mid-land Naturalist.
was a member of the New England Botanical Club, American Association for the
Advancement of Science, Botanical Society of America, American Association
of University Professors, American Biological and Lichenological Society,
American Society for Plant Taxonomy, Sociedad Botanical de Mexico, Illinois
State Academy of Science, American Fern Society, Sigma Xi, California Botanical
Society, International Association for Plant Taxonomy, and Indiana Academy
Clifford Petry 1887-1970
death of Loren Clifford Perry, professor of botany, emeritus, on May 4 ended
the career of an outstanding student and teacher of botany. Not only was Dr.
Petry renowned as a teacher of introductory botany to thousands of students
at Cornell but his dedication and vitality became equally famed during his
retirement years on Cape Cod.
was born of Quaker parentage in New Paris, Ohio. His first interest was engineering
and his B.S. from Earlham College in 1907 was taken in that field. A year
later he earned a second B.S. at Haverford College. During the following two
years he taught science in the high school at Urbana, Ohio. On the advice
of another noted teacher of botany, Millard Markle of Earlham, he decided
to 'further his interest in plants by pursuing graduate studies in botany
at the University of Chicago. There he earned the M.S. in 1911 and the Ph.D.
first post was an Instructorship at Syracuse in 1914. He was raised to Assistant
Professor in 1916 and Associate Professor in 1919. From 1922 to 1924 he was
on leave from Syracuse as Acting Assistant Professor of Botany at Cornell.
During the years 1919 to 1925 he served as Director of Summer Session at Syracuse.
In 1925, he was appointed Professor of Botany at Cornell where he remained
until his retirement in 1955.
valuable was his effectiveness in the teaching of introductory botany. An
inveterate traveller and observer of all aspects of the environment, he felt
that facts should be used as tools in the solution of problems rather than
as things to be memorized and stored away. As a result, his lectures emphasized
ways of learning and using rather than memorizing. His students were encouraged
to think. Two tangible results of this attitude were a laboratory manual for
introductory botany written in collaboration with E. M. Palmquist and a "Keys
to Spring Plants" written in collaboration with W. C. Muenscher. Approximately
12,000 students were exposed to his botanical wisdom during the 30-year span
of his teaching at Cornell.
research interest centered around the first plants to occupy dry land (Devonian
period). His early collecting expeditions to the Gaspe Peninsula resulted
in a splendid collection of Devonian plants at Cornell that has been increased
by his students until it is now one of the best in the world. His enthusiasm
led others to this area of study and the plants of this period have now become
a critical item in studies of plant evolution. Petry's enthusiasm for paleobotany
also led him and Ralph Chaney of Berkeley to found in the Iate thirties a
paleobotanical section of the Botanical Society of America. Petry served as
the second chairman of this group (1938).
Petry served botany also through participation in its national organization,
the Botanical Society of America. In 1933 he was elected to be its Secretary
for a three-year term. In 1937 he was elected Vice-President of the Society.
From 1936-1939 he was a member of the Society's Committee on Education where
he was instrumental in the production of two publications "An Exploratory
Study of the Teaching of Botany in the Colleges and Universities of the United
States" and "Achievement Tests in Relation to Teaching Objectives in General
College Botany." Both of these reflected some of his own philosophy in teaching.
extent of Dr. Petry's dedication to observation, to interpretation, and to
teaching can be measured by his activity in the fifteen-year period following
his retirement in 1955.
taught for various periods at the University of Missouri, Hofstra College,
University of Utah, and Welles-Iey College. He was also a member of a national
panel of lecturers sponsored by the American Institute of Biological Sciences.
In 1960 he moved permanently to Cape Cod and continued to lecture locally
and abroad, one series at The University, Reading, England.
Cape Cod he became associated with the Cape Cod Museum of Natural History,
Brewster, Massachusetts for which he led trips for young and old to view geological
phenomena, salt marsh plants, and wildlife in general. He cut and marked nature
trails. He lectured on a range of topics from edible mushrooms to salt marsh
glaciation of New England. He devised exhibits and experiments to attract
and to explain.
was no surprise that in 1966 his Alma Mater, Earl-ham College, rewarded him
with an honorary D.Sc. nor that he was immensely pleased to be presented for
that degree by the late Millard Markle who had originally influenced him to
go on to graduate work at Chicago.
R. H., AND R. A. SCOTT (Editors). Aspects of Palynology. John Wiley &
Sons, Inc., New York, 1969. vii + 510 pages. $24.95.
science of pollen analysis is relatively new, yet it already has achieved
an important role in the study of such diverse disciplines—to mention
only a few—as paleoecology, archaeology, paleobotany, and petroleum
geology. During the past several decades the recognized importance and application
of palynological research has resulted in a large body of published data,
little of which appears in the form of syntheses or comprehensive reports.
As stated by the editors, R. H. Tschudy and R. A. Scott, Aspects of Palynology
is designed to provide just such syntheses to the beginning palynology student,
teacher, and researcher through a collection of articles on the nature, scope
and applications of the study of fossil pollen and spores.
book presents a collaborative effort by 14 separate authors. The first five
chapters provide a general background of data essential to the understanding
and study of fossil pollen and spores. Represented in these chapters is an
introduction, followed by discussions of pollen/spore classification and descriptive
teminology by R. H. Tschudy, pollen ontogeny and pollen wall development by
A. O. Dahl, systematic and nomenclature guide-lines used in palynology by
J. M. Schopf, and a section on the relationship of sedimentation to palynomorph
preservation by R. H. Tschudy. The next two chapters of the book cover subjects
pertaining to pollen analytical interpretations. The first of these, by J.
W. Funkhouscr, examines the reliability of pollen samples and provides a useful
summary of frequent sources of contamination during the collection and subsequent
pollen analysis of sediments. The second of these two chapters, by R. H. Tschudy,
is devoted to a summary of the major concepts around which pollen analytical
interpretations are based. The remainder of the book, except for one chapter
by W. R. Evitt on non-palynomorphs often encountered in pollen preparations,
contains a series of chapters devoted to a brief summary of palynomorph assemblages
through geologic time. These include discussions of Precambrian and Early
Paleozoic palynomorphs by J. M. Schopf, Devonian spores by J. B. Richardson,
Mississippian and Pennsylvanian microfossils by R. M. Kosanke, Permian palynomorphs
by G. F. Harr, Triassic spores and pollen by W. G. Chaloner, Jurassic and
Early Cretaceous micro-fossils by N. F. Hughes, Late Cretaceous and Early
Tertiary palynomorphs by J. S. Penny, and Late Cenozoic palynology by E. B.
for its expense, Aspects of Palynology is well suited for use as a college
textbook or as a reference source. The chapters are arranged in a logical
chronological sequence, and in most cases they are well illustrated (except
on page 352 where part of one illustration is faded and illegible) and contain
sufficient bibliographies. However, there are a few disconcerting instances
in which important literature either was not mentioned or was cited incorrectly.
In chapters dealing with the relation-ships of palynomorphs to sedimentation
and factors that affect reliability of samples, neither of the authors (R.
H. Tschudy nor J. W. Funkhouser) cites or mentions the contributions of G.
W. Dimbleby (1957) on pollen preservation and palynomorph movement in terrestrial
soils, S. Goldstein (1960) and W. C. Elsik (1966) on biologic degradation
of pollen and spores, A. G. Sangster and H. M. Dale (1961, 1964) on differential
pollen preservation in lacustrine and bog deposits, H. Tauber (1965) on differential
pollen dispersion, and E. J. Cushing (1967) on differential pollen preservation.
In several instances references cited in the text either do not agree with,
or are omitted from, the references cited in the bibliography. Some examples
include: references cited on page 35 include (Michel, 1940) and (Freytag,
1954) yet in the bibliography they appear as (Michel, 1950) and Freytag, 1964)
; on page 141 a reference is listed as (Gothan and Weyland, 1964), yet in
the bibliography it is listed as (Gothan and Weyland, 1954); on page 319 a
reference is listed as (Seward, 1904) yet it does not appear in the bibliography;
and on page 326 (Erdtman, 1948) is cited, yet omitted from the bibliography.
than three-fourths of this book is devoted primarily to the pollen record
of the Pre-Quaternary portion of the geologic record and thus will undoubtedly
and logically be utilized most widely for studies of Pre-Quaternary palynology.
Yet this book should also be of interest to Quaternary palynologists since
there are useful chapters on applied palynology, pollen ontogeny, Late Cenozoic
palynology, and non-palynomorph microfossils encountered in palynological
samples. The merits of this book outweigh its minor faults and in spite of
its initial expense, it should provide a useful reference source for many
M. Bryant, Jr.
of Pesticides on Fruit and Vegetable Physiology. National Academy of Sciences,
Principles of Plant and Animal Pest Control. Vol. 6. Washington, D.C. 1968.
90 pages. $3.25 + .20 handling.
volume does an excellent job of bringing the botanist into contact with literature
on growth regulators in a field esoteric even to most physiologists. In nearly
every chapter, the authors conclude that there is an acute need for more careful
and thorough investigation. Those who look for an understanding of mode of
action will be disappointed. Rather, this is a compendium of who, what, when,
and where with relatively little "how." Fruits obviously offer some unique
opportunities for research with aging and senescence at least partially delimited
by the climacteric. The book does a good job, so far as I can tell, of discussing
current research and problems in this interesting area.
HYAMS, E. AND W. MACQUITTY. Great Botanical Gardens of the World. Macmillan,
New York, 1969. 288 pages. S35.00.
like the reviewer, you have been delighted by visits to some of the world's
leading botanical gardens, perusal of this magnificent volume will bring back
many pleasant memories as well as to whet your appetite for travel to see
gardens you may have missed. In his foreword to this volume, Sir George Taylor,
Director of the Royal Botanic Gardens at Kew, has written. "but no work known
to me deals with the major botanical gardens of the world with the scope and
on the lavish scale of Great Botanical Gardens of the World. The author, Edward
Hyams, and the photographer, William MacQuitty, have chosen a broad canvas
on which to present their facts and impressions of their gardening odyssey."
Of the 42 botanical gardens listed by name in the table of contents, 18 are
in Europe, 8 in North America, 5 in the USSR, 6 in the "tropics," and 5 are
listed as "southern hemisphere." A separate section is devoted to several
gardens in japan. At the end of the book, and after the detailed index, are
five pages on which 525 of the world's botanical gardens, compiled from the
International Directory of Botanical Gardens (Utrecht, 1963), arc identified
and located by corresponding numbers on accompanying maps. A minor error involves
a picture on page 97, labeled as a view of SchOnbrunn park, but actually a
view at Belvedere park, also in Vienna. Botanist-gardners who are also interested
in photography will be pleased to note the Technical Data on cameras, lenses,
etc. (page 13). In addition to the many excellent black-and-white photographs,
64 of the pages are in brilliantly reproduced color.
M. AND S. ASANO. Biological Flora of Japan, Vol, 1. Sympetalae—1 -f-
Introductory Volume. Tsukyi Shokan Publ. Co. Ltd. Tokyo, 1969.
work is an ambitious undertaking to represent by line drawing, photograph,
and terse statements the ecological status of (all?) the native and introduced
plants of Japan. Volume I (of a projected five volumes) covers largely composites
and a few others of the Sympetalae. The basic format of the first volume is
a devotion of two pages to each species selected by the authors. The lefthand
side consists of terse statements in Japanese and English on habitat including
distribution, climate, soil, physiography, and vegetation type; and then life-form
including dormancy, dispersal unit, and other ecological features, together
with a photograph of the plant in some habitat situation. The righthand side
of the pair of pages illustrates in excellent line drawings the plant in question.
The first volume was distributed in a handsome box together with a hardbound
volume of introduction. The introduction is also bilingual and briefly explains
the in-tent of the work and the bases on which the terse eco-Iogical notes
are made. As a work of art, the present volumes represent a real triumph.
Both the photographs and the drawings are well executed, although I am taken
a bit aback at the rather overpowering exhibitions of the underground portions
of a plant—masses of roots which don't seem to tell too much.
when I ask the question, "What scientific function does this work fill?" I
am less easily convinced of its great worth and significance. To my mind a
biological flora is one in which a great deal of ecological life history data
are accumulated for each taxon in a regional flora. The model for such biological
flora is probably the long-term publication of ecological life histories in
the British Journal of Ecology. The present work does not measure up in terms
of in-depth studies to this extent. Habitat features and life-form descriptions
are so terse as to be much too general. However, the work should be valuable
as a guide to ecological interpretations of the Japanese flora. Certainly
plant ecologists working on specific communities would want to compare their
own evaluations with the summary notations given in the present work. And
as a device for corroborating identification, the work should be eminently
useful, especially with the habitat photographs and the derailed line drawings.
primary value of this series to botanists other than those in Japan would
be: (1) to have a pictorial reference for validating plants collected in Japan
or known to be of Japanese origin; and (2) to serve as a model with modifications
for other so-called biological floras whose intent is to describe the habitat
features of the members of the regional flora.
P. H. Flora of Turkey and the East Aegean Islands. Edited by P. H. Davis,
assisted by D. P. Chamberlain and Victoria A. Matthews, University of Edinburgh.
Volume three, Edinburgh, at the University Press, 1970; xvii, 628 pages, 15
full-page line illustrations, 98 maps. Price 9 pounds, 9 shillings. North
American agent, Aldine Publishing Company, 526 South Wabash Avenue, Chicago
60605. U.S. Price $33.50.
volume, devoted entirely to the large family Leguminosae, has been produced
with the high standards set in Volumes 1 and 2, the latter of which was reviewed
in this Bulletin in December 1968. A simplified terminology for summarizing
internal distribution, similar to that used in Volume 1, has been adopted
in preference to that of using solely the names of the Turkish vilayets. Contrary
to earlier practice, new scientific names and new combinations are published
in the text. Much of the text (including the largest genus, Astragalus, with
372 species) has been contributed by the research assistants, Dr. Chamberlain
and Miss Matthews. As in earlier parts of the flora, treatments of certain
genera or parts of genera have been contributed by specialists in these groups.
Leguminosac is a very large and diverse family in this part of the world,
and American botanists will find much of interest in the genera that are known
chiefly because of a few introduced species. The number of native and introduced
genera in the Turkish flora is not particularly large (68), but several genera
are rep-resented by numerous species, as Trifolium with about 100, Vicia and
Lathyrus with almost 60 each, Trigonella with 49, Onobrychis 46 and Medicago
contents of Volumes 4 and 5 have been announced by the Editor, and it is hoped
that their production can be achieved as promptly as that of the present installment
in this very useful and important series.