238 CRC Handbook of Flowering

CHRYSANTHEMUM MORIFOLIUM

En. Chrysanthemum; Fr. Chrysantheme; Ge. Chrysantheme; Sp. Crisantemo

Kenneth E. Cockshull

INTRODUCTION

The chrysanthemum has been cultivated for more than 1400 years in China and for at
least 1200 years in Japan, 1 and is now one of the most popular cut flowers in North America,
Western Europe, and Japan. Chrysanthemum morifolium Ramat. is a complex hybrid and
its popularity and long history of cultivation have led to the introduction of many thousands
of cultivars showing great diversity of flower form and plant habit. It has recently been
suggested that C. morifolium should be renamed Dendranthema morifolium (Ramat.) Tzvelev.2
Both genera are members of the family Compositae.
Chrysanthemums are mainly herbaceous perennials with lobed leaves, spirally arranged
on erect, branched stems and they bear their flowers in capitula. They perennate naturally
by means of stolons, often terminating in a rosette of leaves, and are readily propagated
asexually by means of stem cuttings taken from basal shoots that develop in the spring.
These are then allowed to flower in their natural season and the cycle repeated the following
year.
The flowering time of most cvs can be controlled by modifying the natural day length.
This has encouraged the development of techniques for producing flowers all year round
(AYR) 3 - 4 which, in turn, has contributed greatly to the present popularity of the chrysan-
themum, cvs selected for AYR production are maintained as vegetative stock plants in LD;
stem cuttings are taken from these, rooted, flowered in SD, then replaced by fresh cuttings.

FLORAL MORPHOLOGY

The individual flowers or florets are small and relatively insignificant, but they are collected
together into heads (capitula) to make showy inflorescences. Each capitulum is surrounded
by an involucre of green bracts, within which the florets are mounted on an enlarged
receptacle. In other members of the Compositae, the florets appear in the axils of receptacular
bracts or scales, but these structures are normally absent from the receptacles of most
chrysanthemum cvs, unless the early stages of inflorescence formation occur in LD. S - 6
There are basically two types of floret. The marginal, ray florets are female and in their
typical form have a corolla that flattens out above a tubular base, to create a long, strap-
shaped tongue or "ligule", hence the term ligulate floret. The central, disc florets are
hermaphrodite and in their typical form have a small, tubular corolla terminating in five
distinct teeth. Modifications to these basic forms have occurred during cultivation and are
sometimes used for classifying flower types, e.g., anemones, quills, spoons, spiders, etc.1-4
Both types of floret have an inferior ovary containing a single ovule.
The proportion of ray and disc florets in each capitulum varies between cvs; "double"
inflorescences have a majority of ray florets, while disc florets predominate in "single"
inflorescences. The proportions are also affected by daylength, 79 by changing the night
temperature, 9 - 10 by the application of growth regulators," l 2 and by the position of the
inflorescence for the terminal capitulum tends to have a greater proportion of disc florets
than the axillary ones. 13 The total number of florets per capitulum varies between cvs,8 but
can generally be counted in hundreds. The number of ray florets is increased by higher daily
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light integrals, 14 higher temperatures, 9JS LD,* and by intercalating some LD during the
period of floret initiation. 7 - 9 -"' Conversely, the number of disc florets is usually increased
by shorter day lengths, 8 - 9 though more were obtained when transfer to LD followed just 7
SD. 7 Transfers to low daily light integrals reduced the number of ray florets formed but
only when transfer was made at the onset of floret initiation, about 14 days after the start
of SD. 17
Each inflorescence is a raceme with the oldest florets around its margin, though the
flowering shoot as a whole is a cyme with the oldest inflorescence at its center, terminating
the growth of the main shoot. The axillary apical meristems are subject to apical dominance
until the terminal apical meristem is removed or initiates an inflorescence; release from
apical dominance then proceeds basipetally. 5 - 18 Thus, if the axillary meristems were also
incompetent to respond to the floral stimulus while subject to apical dominance, this would
have the effect of generating the cymose form of the flowering shoot. 14
Although botanically the chrysanthemum flower is a floret, the individual capitula are
commonly referred to as flowers in horticultural texts. It is in this sense that the term flower
will be used in subsequent sections.
The chrysanthemum is marketed as a cut flower in either "standard" or "spray" form.
The former consists of a stem from which all but the terminal flower has been removed,
while in the latter case, the lateral flowers are retained and the terminal is removed. The
unwanted flower buds must be removed as early as possible so as to increase the dry weight
and quality of those that remain. 19 When producing cut flowers in their natural season, it
is common practice to remove the shoot tip from the rooted cutting ("pinching" or "stop-
ping") so as to produce lateral shoots, of which three are usually retained to form flowering
stems.

APICAL MORPHOLOGY

Popham and Chan 20 2 I described and illustrated the anatomy and morphology of vegetative
and reproductive shoot apical meristems. The morphological changes (Figure 1) form the
basis of an arbitrary scale of reproductive development22 which has been widely used to
quantify the flowering responses of chrysanthemum. The vegetative apical meristem can
grow to 250 p,m in diameter, 6 - 2 "- 23 is slightly domed, and is surrounded by from 9 to 13 or
more leaf primordia and young leaves which form an apical bud. 24 The rate of formation
of leaf primordia is accelerated by both increasing irradiances and air temperatures. 25
The initiation of flower formation is marked by the enlargement of the apical dome to
form a receptacle and by the formation of primordia which retain their entire margins and
grow more slowly than the leaf primordia, though they are initiated more rapidly than
them." 21 These become the involucral bracts which enclose the receptacle and floret pri-
mordia. Floret initiation begins around the outer edge of the receptacle and proceeds ac-
ropetally until the entire receptacle is covered. The timing of these events is influenced by
environmental conditions. Under favorable conditions, rapid enlargement of the apical dome
begins after 7 to 9 SD; floret initiation begins between the 12th and 16th SD and is complete
by about the 24th to 30th SD. 14 -' 6 - 18 - 21 - 26 Because there may be at least 13 leaf primordia
within the apical bud at the onset of flower initiation, it takes from 24 to 30 days for these
to unfurl and reveal the flower bud. The diameter of the receptacle changes dramatically
during this initial period of SD, 6 -"- 21 - 23 and can increase from 240 to 2800 |xm in the course
of 30 SD (cv Bittersweet ). 21
Horridge and CockshulP showed that reproductive development in 'Polaris' was correlated
with a growth in apex size such that the onset of floret initiation, for example, was associated
with a narrow range of apical volumes in a number of different environments. Furthermore,
there appeared to be a critical size of apical dome, below which only leaf initiation occurred
240 CRC Handbook of Flowering

FIGURK 1. Stages in the initiation of the chrysanthemum inflorescence. A: Plan view of an apical meristem at
the transition to inflorescence initiation. The oldest, intact priniordium has a dentate margin and will become a
leaf ( x 150). B: Plan view of a reproductive apical meristem surrounded by bract primordia with entire margins.
The meristem has enlarged considerably and is forming the receptacle ( X 150). C: The receptacle has continued
to enlarge and is forming florets on its flanks. The older bracts which overarch the receptacle have been removed
to reveal the florets ( X 112.5). D: Floret primordia have been formed over almost the whole surface of the
receptacle. The older, outer primordia are clearly differentiating into ligulate ray florets ( x 37.5). Figure A. cv
Miros; Figures B to D. cv Dark Westland. (Figure I A reproduced from Larsen, R., Swedish J. Agric. Res.. 12.
95—102. 1982. With permission. Figures I B to I D reproduced by courtesy of Rolf Larsen. The Swedish University
of Agricultural Sciences, Alnarp, Sweden.)

and above which bract and receptacle formation began. A similar relationship was found in
'Bittersweet'. 27
In certain conditions, e.g., natural LD, the shoot apical meristem may form a flower bud
which fails to grow rapidly. It may have enlarged involucral bracts and is normally surrounded
by leafy lateral shoots arising from the axils of the upper leaves. These buds are often
referred to as "crown" or "break" buds, especially in the earlier literature. Dissection of
them reveals that floret formation is incomplete and at least the central area of the receptacle,
and sometimes its entire surface, is free of florets. The buds are formed in the same way
as normal flower buds; 18 - 21 they have the potential to develop normally but their development
has been retarded or arrested by the environmental conditions in which they are growing. 5 '"

GENETIC INFORMATION RELATED TO FLOWERING

The chrysanthemum is an outbreeding, hexaploid (2n = 6x = 54) but the chromosome

number is variable. 28 There is little genetic information about any character other than flower
color; the inheritance patterns are in doubt, and many cvs are periclinal chimeras. 29 Although
individual cvs flower at a specific time each year, there is continuous variation in the time
of flowering of different cvs. The range extends from June to December under natural
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FIGURE I B

FIGl'Ri: 1C

conditions in Japan and England. 8 - 1 " It appears to be dependent mainly upon two characters,
MY/.., the duration of vegetative growth before flower bud formation and the rapidity of
flower bud development, and their interaction with daylength and temperature. 3 " Shoots both
242 CRC Handbook of Flowering

FIGURE I D

of early-flowering (summer) and late- flowering (autumn) cvs eventually form flower buds
in LD, but they are generally formed sooner and develop more rapidly in SO.8-30-1'
The unselected progenies of crosses between early- and late-flowering cvs showed con-
tinuous variation both for the time of flower bud formation in LD, as assessed by the number
of leaves formed before the bud (LD leaf number), and the speed of bud development in
LD. The two characters were positively correlated, however, so that clones which showed
rapid flower bud development in LD also tended to form few leaves in LD.-12
Recently, more interest has been shown in the genetic variation in LD leaf number, mainly
because of its relevance to "premature budding" and compound spray formation in cut-
flower production.24 A test procedure has been devised to screen clones for this character.11
Data are now available for many of the cvs grown in England,14 and information is being
gained on the inheritance of this character. 35

JUVENILITY AND PLANT AGE

Chrysanthemum seedlings exhibit a phase of juvenility during which they continue to

produce leaves, even when growing in conditions that are normally conducive to rapid flower
formation.-16 This phase can last for many weeks in some seedlings36 and it appears to be a
property of the shoot tip, for shoots from axillary buds on the seedling axis form flowers
more readily. 37 After flowering, the chrysanthemum can be vegetatively propagated by means
of stem cuttings and similar periods of unresponsiveness can occur in this material which
is no longer juvenile.
In cvs that require vernalization, the unvernalized state (see next section) represents a
phase of unresponsiveness which has similarities with juvenility in seedlings; both are
properties of the shoot tip, 17 - 3 * and are less evident in shoots arising from axillary buds on
the main axis. 3719 In material that does not require vernalization, axillary meristems released
from apical dominance by decapitation of the main shoot may be temporarily unresponsive
 Volume II 243

FIGURE 2. The relationship between the time of receptacle initiation (apical stage 2)~ in lateral shoots
of chrysanthemum cv Bright Golden Anne, and plant dry weight, when grown in SD (8 hr) at irradiances
of 10.9 (•),21.7 (A), 43.4 (O), and 86.8 W/m2 (A) combined factorially with three CO, concentrations
(not separately identified). The plants were pinched 1 week after the start of SD.45

if transferred to SD at the time of the decapitation. 40 The duration of this period, as assessed
by the number of leaves produced, varies with genotype.8-41
After this initial unresponsive phase, it appears that increasing shoot age does not greatly
affect the flowering response in SD, 42 - 43 although small plants can remain unresponsive for
some time.44 This may be associated with the need to reach a minimum plant dry weight
(Figure 2). 45 - 46 Age has a profound influence in LD, for rooted cuttings form flower buds
progressively earlier the older the shoot is from which the cutting is taken. 47 The basis for
this response is discussed later, but it is not unlike a physiological aging of the apical
meristem which eventually results in flower bud formation after the initiation of a maximum
number of leaves characteristic of each cv. 48 The process is apparently unaffected by re-
moving and rooting the upper part of the shoot to form a cutting. 47 -' 66

VERNALIZATION

Cuttings made from the basal shoots of certain cvs initiate flowers sooner if they are
exposed to low temperatures for a few weeks prior to exposing them to SD. 4y - 50 The re-
quirement for vernalization appears to have been eliminated from cvs selected for year-round
flower production, 1 though this may reflect the fact that they are usually propagated from
lateral shoots rather than basal ones. 51 It is also absent from many cvs that are propagated
in the spring each year from basal cuttings. 50
The literature on the subject has been well reviewed, 1 5 2 5 3 but briefly, basal shoots must
be exposed to low temperatures for at least 3 weeks to obtain the maximum acceleration of
flowering.49 Treatment can be given while the shoots are still attached to the parent plant
or after they have been removed and rooted as cuttings.54 A temperature of about 4°C was
more effective than one of 10°CS4 and it was effective when given only at night. 4y The
244 CRC Handbook of Flowering

chilling requirement can normally be met, therefore, by overwintering stock plants or cuttings
either in the field, in cold frames, or in cool glasshouses. 49 52 A short period of growth in
LD after vernalization accelerated flower initiation in subsequent SD.55
The low temperature is perceived by the growing point at the tip of the main shoot31* and
by meristems in the leaf axils and at the base of the plant. 51 The basal growing points, which
normally remain dormant until the plant has flowered, slowly lose their vernalized state as
the plant ages with the result that they have to be revernalized at the end of the growing
season. 49 - 51 The other growing points that were present during the original chilling treatment
retain their vernalized state and are able to pass it on to further growing points as they are
formed.3"
A vernalized shoot can be devernalized if exposed to a period of high temperatures (>24°C)
in low-light conditions,5'1 but only if this treatment is given within 3 weeks of vernalization. 51
The response suggests that carbohydrate status may be involved, yet defoliation in full light
does not cause devernalization and feeding sugars to plants during devernalization cannot
stop the process.5fi A devernalized shoot can be revernalized by exposure to a second chilling
treatment. 56 Vernalization enables shoots to form flower buds both in SD and in LD, although
they may not develop to anthesis in the latter, whereas unvernalized shoots may remain
vegetative for over a year in LD.49 The unvernalized and devernalized states appear to be
equivalent, and both result in the production of leafy rosettes, especially when plants are
grown in SD at 16°C in low light. 51
Although the vernalized and unvernalized states appear to be stable, at least when plants
are grown in full light at 16°C, the stability is a property mainly of the original shoot tip.
In vernalized shoots, basal meristems become devernalized in time, 51 while in unvernalized
shoots the upper, axillary meristems on old plants appear to become vernalized. 3 '' This
behavior suggests that interactions within the plant may also affect the ability of meristems
to respond to flowering stimuli in a manner similar to their influence on juvenility (see
above).

PHOTOPERIODIC RESPONSE

Individual cvs flower at the same time each year but the range of flowering times extends
from early summer to winter in Europe and North America and can encompass the whole
year with Japanese cvs grown under protection. 57 The autumn- and winter-flowering types
are SDP.' cvs selected for AYR flower production by daylength control in heated glass-
houses are usually chosen from this group and are classified in terms of their speed of
flowering in SD. The 10-week response group signifies that its members will reach anthesis
in 10 weeks from the start of SD treatment at 16°C under autumn conditions. 3 Members of
the 14-week response group reach anthesis in 14 weeks of SD and will flower later, therefore,
than those of the 10-week response group when grown under natural conditions. Summer-
flowering types, by definition, must reach anthesis in LD and so have been regarded as
DNP.44 More recent investigations have shown that most initiate and develop flowers more
quickly in SD8 30 - 31 and so their flowering time could also be modified, if desired.5*
Flower initiation occurs eventually in all cvs when grown in LD 1 6 8 2 4 4 K 5 2 though this
may take many months in unvernalized shoots of cold-requiring cvs,"' 49 - 59 but it generally
occurs earlier when they are grown in SD. Initiation is, therefore, a quantitative response
to SD. Flower bud development is also a quantitative SD response in early-flowering cvs
but a qualitative response in late-flowering ones as they are unable to develop their buds to
anthesis in LD. 5 60
Flower initiation in LD can be regarded as resulting from an autonomous induction process
which proceeds independently of any external signal. It is related in some way to activity
within each apical bud, possibly to the attainment of a critical size of apical meristern, 23 - 61
 Volume II 245

for the removal of expanded leaves has no influence 62 and adjacent axillary mcristems can
remain vegetative, hence the formation of "crown" buds. These adjacent meristems, how-
ever, do form fewer leaves than the main shoot, indicating that there has been some carry-
over of induction/ It also appears that the shoot tip can be removed and rerooted without
influencing the process of autonomous induction. 4 7 For this reason it is desirable to study
the response in axillary meristems recently released from apical dominance. 4 " It can be
quantified in terms of the number of leaves initiated by an apical mcristem prior to flower
initiation."- 48 This number is stable under constant conditions 25 but shows great genetic
variation. LD leaf numbers ranging from 20 to 94 have been recorded from different cvs
grown under summer LD conditions in the U.K. 1 4 Leaf number is influenced by the envi-
ronment (see below).
Photoperiodic induction does require the presence of expanded leaves, 1 - 41 though one leaf
can be sufficient; 6 ' 7 or 8 SD in succession are as effective as continuous SD in inducing
flower initiation, for similar numbers of leaves are formed before the terminal flower bud. l4 - 64
This is to be expected, as the first signs of flower initiation can be detected at the terminal
apical meristem within 8 SD 14 -" 1 — 4 SD in succession are less effective, but still induce
flower initiation before it occurs in LD, 6567 and at a lower leaf number than in LD. 1 4 They
are even less effective if LD are interspersed among them: 65 2 SD in succession caused bud
formation more than 25 days later, but prior to its occurrence in LD. 67 When fewer than 8
SD are given in succession it is generally only the terminal apical meristem that is committed
to flower initiation, 14 i.e., a "crown" bud is formed. 6567 As more SD are given, so the
number of axillary meristems committed to flower initiation increases in a basipetal
progression. l4 - 67 - 68
The effect of daylength on flower initiation is of the general form shown in Figure 3,
though it has been suggested that it can be separated into two distinct components, a SD
response (12 hr or less) and a LD one (14 hr or more). 69 In Figure 3 there appears to be no
critical daylength but a range of daylengths over which there is a gradual change from rapid
to delayed flower initiation. The range over which this occurs varies with the cv. It begins
at a daylength of about 13 hr for cvs in the 10-week response group grown at 16°C.™ 7: In
general, the longer the response group the shorter the maximum daylength at which rapid
flower initiation still occurs. 7 "- 71 Flower initiation is inhibited at daylengths of only 2 hr."
The effect of daylength on flower bud development is more complex. The buds of some
early-flowering cvs will reach anthesis in continuous light, 72 but most develop more rapidly
in shorter day lengths. 8 - 10 - 31 - 72 The flower buds of late-flowering cvs will not normally develop
to anthesis in LD, 5 - 8 - 60 - 72 and the maximum daylength at which development will proceed
is usually shorter than that at which rapid flower initiation occurs.70 7 2 7 4 The inhibitory effect
of LDon flower development is dependent upon the time of treatment. When flower initiation
has occurred in LD, development is arrested in continuing LD unless the lateral, vegetative
shoots around the terminal flower bud are removed or the bud itself is removed and rooted
in LD.5 If bud formation is induced by exposure to 14 SD, the lateral shoots also form buds
and the terminal will develop slowly to anthesis in summer LD.32 After about 28 SD, the
terminal bud will develop to anthesis with virtually no delay, 75 especially if the lateral buds
are removed,64 and later still, e.g., when flower color is visible transfer to LD has no effect
on the development of either the terminal or lateral flower buds. 3 - 4

NIGHT-BREAK LIGHTING

As flower initiation occurs eventually in all daylengths,'- a48 - 52 the inhibitory effect of
different NB treatments can be compared either in terms of the numbers of leaves produced
or of days that elapsed prior to flower initiation. It is then possible to identify treatments
which give maximum inhibition. Unfortunately, most investigations have been terminated
246 CRC Handbook of Flowering

FIGURE 3. The effect of daylength on the inhibition of flower initiation as assessed by the number
of leaves formed prior to the terminal inflorescence, cv Snowdon.

after 2 or 3 weeks, i.e., before flower initiation occurred in all treatments. It is then impossible
to determine whether the vegetative plants would have remained in that condition for another
day or another month.
The inhibition of flower initiation becomes more pronounced as either the duration 22 - 76 7y
or the flux density 77 x2 of the light increases when given as a NB near the middle of the
night. The response is, therefore, a quantitative one. When light from "cool-white" tubular
fluorescent lamps was given as a NB lasting from 1 to 4 min at different illuminances, the
degree of inhibition achieved was the same in all combinations having the same light integral
(e.g., 4.3 klx/min). 78 Thus, the response shows some measure of reciprocity. Cvs differ in
their sensitivity to NB lighting. 22 - 44 - 77 -*"- 8 '- 8 - :i The illuminance that gave maximum inhibition
of flower initiation varied from 90 to 150 Ix, according to cv, when light from incandescent
lamps was given continuously for 5 hr about the middle of a 16-hr night (27 to 45 klx/
min). 80 The amount of radiant energy (400 to 700 nm) received during the day can also
influence the effectiveness of a NB treatment; more energy is required during the NB when
the daily light integral is high. 79
The inhibitory effect of a 1-min NB per night can be detected when it is given at an
adequate illuminance, e.g., 6.46 klx from "cool-white" tubular fluorescent lamps inhibited
flower initiation for 14 days.™ Light from incandescent lamps, at an illuminance of 12.9
klx for 1 min was less effective, although the energy content per lumen in the 600 to 700
nm waveband was higher than that of the fluorescent lamps. 1K Similarly, at a standard
illuminance of 430 Ix, flower initiation by 'Honeysweet' was inhibited for 14 days by nightly
exposure to 12 min of light from "cool-white" fluorescent lamps, but required 48 min of
light from incandescent lamps (i.e., 20.7 klx/min) to give the same degree of inhibition. 77
Incandescent lamps were also less effective than "cool-white" fluorescent lamps when
compared at the same level of installed power.*1 " When compared at equal incident irra-
diances (400 to 700 nm), the emission from the "Gro-lux" fluorescent lamp was more
effective than that from incandescent lamps when given for 30 min each night. 82
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Despite these differences, the incandescent lamp remains the light source most commonly
used in commercial practice, 3 - 4 mainly because of its low capital cost. Integrals of between
24 and 30 klx/min are obtained by lighting for 4 to 5 hr each night in winter at an illuminance
of about 100 lx. 3 - 4
It has been suggested that the relative ineffectiveness of the incandescent lamp for NB
treatments is due to the high FR content of its emission. 77 R, of an unstated irradiance,
inhibited flower initiation for 14 days when given as a NB for 9 min or longer each night.
The effect of R was partial'y reversed by a similar duration of FR light given within about
30 min of the R treatment, but treatment with FR light alone for 81 min also had an inhibitory
effect. 22
Early experimenters began their NB treatments at midnight, 84 and Post85 stressed that the
objective was to ensure that any period of uninterrupted darkness did not exceed 9 hr each
night. Even 10 hr of darkness, either following a 6-hr daylength extension with incandescent
lamps (215 lx), 86 or preceding one using "cool-white" fluorescent lamps (19.4 klx)7* failed
to cause rapid flower initiation. Furthermore, light from "cool-white" fluorescent lamps
for 1 min at 6.5 klx was inhibitory when given at any time from the 6th to the 9th hr of a
15-hr dark period, and at 19.4 klx was inhibitory from the 3rd to the 9th hr. 78 Treatment
after the 3rd hr of darkness would have been followed by almost 12 hr of darkness. In
commercial practice, 4 or 5 hr NB are usually placed around midnight so as to divide the
night into equal portions.3-4 There seems no reason, however, why they could not begin at
midnight, if this takes advantage of lower electricity tariffs, and if the period of darkness
before midnight does not exceed 9 hr.
When light was given throughout a 16-hr night, using five different light sources to
provide a low illuminance of 10.8 Ix (10.4 klx/min), incandescent and high-pressure sodium
lamps delayed flowering more effectively than either "cool-white" fluorescent, metal halide,
or high-pressure mercury lamps.87 Essentially similar results were obtained with higher
illuminance levels from incandescent and fluorescent lamps.88 It was suggested that the high
energy content in the 700 to 850 nm waveband of the emission from the incandescent lamp
contributed to its effectiveness when used in this way, 87 although this same waveband is
thought to make it less effective when used as a NB light source.77
Light that was deficient in FR wavelengths effectively inhibited flower initiation when
given for 8 hr before an 8-hr day, but not when given after. Light from incandescent lamps
was effective in both periods.89 Short periods of FR light at the end of an 8-hr day have no
effect on flower initiation, 73/JO though the effectiveness of a subsequent, short R NB is greatly
enhanced by this pretreatment.90 The data suggest that the presence of a high level of
phytochrome in the Pfr form at the start of the dark period promotes flowering or else inhibits
the production of an inhibitor of flowering, while the presence of Pfr later in the night
inhibits flowering. 73
Reciprocity does not hold when a given amount of light from incandescent lamps is spread
over more than one light period each night. Thus, a total of 5.2 klx/min was relatively
ineffective in inhibiting flower initiation when given in one 12-min period, but when this
same total was given over two 6-min light periods separated by 40 to 60 min of darkness
it became very effective. 76 The inhibitory effect was less when the intervening dark period
was shorter than 40 min. When 12 min of light, providing a lower integral of 2.6 klx/min,
was broken up into many short light periods distributed over 4 hr in the middle of the dark
period, the degree of inhibition now increased as the dark interval separating successive
light periods was reduced to 30 min or less.7'1 Cathey and Borthwick called this pattern of
lighting, "cyclic lighting". 76
In general, the inhibitory effect of cyclic lighting, using incandescent lamps, increases as
the dark duration per cycle is reduced, 76 - 7791 and as either the illuminance 767791 92 or the
duration of the light period in each cycle increases.76 9I - 92 For these reasons, the most effective
248 CRC Handbook of Flowering

way of using light of a given illuminance is to provide it continuously throughout the duration
of the NB period. 81 ' 9! In commerce, however, the need is to find the most effective way of
using a given quantity of electrical energy. Since cyclic lighting greatly enhances the in-
hibitory effect of a given light integral and gives adequate control of flowering for the
cultivation of most cvs, it is a technique that is widely used in commercial cropping. In
practice, cycles of 1 min of light in 5 min, 6 in 30, 10 in 30, and 15 in 30 are commonly
used. '-4 The first two would normally be recommended only when an illuminance of at least
110 Ix can be achieved.
As with continuous lighting, light from incandescent lamps seems less effective than that
from fluorescent lamps, 77 - 82 - 88 but cyclic lighting did not appear to enhance the effect of a
given quantity of radiant energy from fluorescent lamps."2 If confirmed, it would suggest
that no advantage would be gained by using cyclic lighting with fluorescent lamps. Fur-
thermore, whereas light from incandescent lamps became more effective when it was supplied
cyclically throughout a 16-hr dark period, that from fluorescent lamps became less effective. Hx
Full inhibition of flower initiation was restored if cyclic lighting was given for only 12 of
the 16 hr. 88 The response appears analogous to that obtained with daylength extensions with
light from fluorescent lamps described earlier.
Flower development appeared to have more stringent requirements than flower initiation,
for when treatments that failed to give full inhibition of flower initiation were prolonged,
they still arrested flower development. 91

TEMPERATURE

As night length has such a profound influence on flower development, it is, perhaps, not
surprising to find that many investigations have shown that the temperature of the night
period has more influence on these processes than that of the day.' This has been interpreted
as evidence that flowering is dependent upon reactions that are specific to the dark period.
More recently, however, it has been shown that many cvs will initiate and develop flowers
as rapidly in night temperatures of 10°C as 20°C, provided that the night and day periods
are of equal length and that the day temperature is 20°C in the former case and 10°C in the
latter.1'1 These data suggest that flowering is equally responsive to the day and night tem-
peratures and that the mean temperature over the whole daily cycle determines the time of
flowering. Thus, night temperature appeared to be of especial importance in the earlier
investigations because it was always given for a much longer period than that of the day
and, therefore, contributed more to the mean 24-hr temperature. Other recent investigations
have concentrated on reducing the night temperature for part of the night94 96 or part of the
SD period.97-98 In general, these treatments delayed flowering, as would be expected from
their effect on the mean 24-hr temperature.
Flower initiation in SD occurs more rapidly as the temperature is raised from 10°C to
about 25°C in most cvs and will occur at higher temperatures. 1 - 99 It shows a quantitative
response to temperature as it does to most other environmental variables. Flower develop-
ment, on the other hand, shows a more qualitative response.100 This is particularly evident
in the group of cvs classified by Cathey 101 as thermonegative, in which flower development
is promoted by increasing temperature up to about 16°C, but is then often arrested at 21°C
or higher. IO(U()I A similar response is seen in his thermopositive and thermozero classes, but
with the optimum shifted to higher temperatures;1 '°° development was delayed at day15 or
night 100 temperatures of about 28°C and completely arrested at an average night temperature
of 32102 to 38°C.91
Cathey's classification of cvs into three response groups; thermopositive (flowering in-
hibited by low temperature), thermonegative (flowering inhibited by high temperature), and
thermozero (flowering not especially sensitive to either high or low temperatures) was based
 Volume II 249

initially upon the effect of temperature in the LD period on the flowering behavior of the
plants when transferred to SD. 101 The flowering of thermozero cvs was relatively unaffected
by the temperature of the LD period, while that of thermopositive cvs was particularly
delayed at 10°C if the LD period was also spent at 10°C.U" There was a carry-over, therefore,
of the effect of low temperature during LD (persisting sometimes after low-temperature
treatment of the stock plants), 103 which has since been noted in other cvs.43-104 The effect
was often characterized by arrested development of the terminal flower bud accompanied
by a delay in the initiation of flowers on the upper side shoots. 43101J(BJ04 In later papers,
the classification was restricted to the responses to temperature in the SD period1 105 and
was used in this form to classify more than 120 cvs. l06 Most cvs used today in AYR flowering
programs would be classified as thermozero on either basis, for they are capable of flowering
in a wide range of temperatures,'"' and show no carry-over effect of spending the LD period
at 10°C l07 or25°C. 108
When two autumn-flowering cvs were grown throughout in LD of continuous light, flower
initiation was delayed at 10 and 28°C relative to 16°C.25 This did not necessarily result in
an increase in the leaf number at the lower temperature because the rate of leaf initiation
was also retarded. Flower development was arrested at all the temperatures used from 10
to 28°C. It was also noted that whereas the axillary meristems below the terminal flower
bud tended to produce leafy shoots at 16°C and above, they rapidly initiated flower buds at
10°C. Flower initiation in the summer-flowering 'Golden Stardust' occurred very rapidly at
a low leaf number when grown at 16°C in continuous light, but was greatly delayed at
10°C.25 Flower development did proceed towards anthesis in this cv, but only at 10 and
16°C and not at 22 and 28°C.2Ii These latter results confirm the view that temperature has
a greater influence than daylength on flower initiation and development in this group of
cvs. 1 - 44

LIGHT INTENSITY

Delayed flowering has frequently been observed under natural low-light conditions 18 - 52 - 104
and early investigations with artificial light showed initiation was greatly delayed when the
"day" period was spent at a low illuminance. 109 The results of a series of investigations in
artificially lit, controlled-environment cabinets, reviewed by Cockshull,14 revealed that flower
initiation in SD occurred earlier the greater the daily integral of radiant energy (400 to 700
nm) that the plants received, in the range 0.31 to 2.50 MJ/m 2 /day. 14J1(1 Furthermore, flower
formation proceeded more rapidly 14 and uniformly 17 "° at the higher light levels and anthesis
was reached earlier.' 4 • no The plants appeared to be most sensitive to light level during the
first 4 weeks of SD. Transfer from a standard environment providing 1.25 MJ/m 2 /day to
one providing only 0.31 MJ/m 2 /day, delayed flower initiation when made for 2 weeks at
the start of SD. As a result, anthesis was also delayed. Transfer for 2 weeks at the start of
the 3rd week of SD delayed completion of the flower bud and also delayed anthesis. Transfer
later, when further floret formation had ceased, had almost no effect on flower development
and anthesis was not delayed. 17
The different daily totals of incident radiant energy were obtained by providing constant
irradiances of 10.9 to 86.6 W/m 2 (400 to 700 nm) for 8 hr each day. When the irradiance
was varied between 11 and 44 W/m 2 or 22 and 87 W/m 2 within each day, the flowering
response was related to the daily total of radiant energy rather than to the maximum irra-
diance." 1 In addition, the plants were capable of integrating the radiant energy receipt over
at least 2 days and then responding accordingly, for the effect of low light in 1 day could
be countered by the provision of high light in the following day. 112 These results suggested
the use of supplementary lighting in winter to promote more rapid and even flowering. 1 ' 3
Irradiance had a marked influence on the autonomous induction of flower initiation in
250 CRC Handbook of Flowering

continuous long days, 25 and the time to flower initiation and the number of leaves formed
prior to the flower bud decreased as the visible irradiance increased from 7.5 to 30 W/m 2
(0.65 to 2.59 MJ-m 2 d ~ ' ) . The number of leaves formed was not further reduced at irra-
diances greater than 30 W/m 2 but the rate of leaf initiation was increased and so flower
initiation occurred even earlier.25 High light levels in LD, prior to SD, gave earlier flowering
in some cvs 165 but not in others." 4

NUTRITION

Although there is variation among cvs in their response to nutrition, the flowering of
many is indifferent to the levels of the major nutrients in the rooting medium, provided
daily integrals of radiant energy are high.3-"5""1* Under low light conditions, however, the
delaying effect of low nutrient levels is more evident, 3 and while high potassium levels then
promote flower initiation," 9 high nitrogen levels delay it." 8 ' 119
Of the minor elements, boron deficiency frequently causes abnormal floret development,
producing tubular or quilled florets,"*-120 l21 incurved inflorescences,118-120 and necrotic areas.123
Iron deficiency delays flowering and can arrest flower bud development," 7 - 118 and copper
deficiency can have similar effects. " 8 - 1 2 1 - 1 2 3 1 2 4 Deficiencies of boron, iron, and copper often
interact" 8 - 121 and the effects of copper deficiency may sometimes be seen only in the presence
of iron deficiency. 124 Severe copper deficiency (<0.0015 ppm Cu in the rooting medium)
delayed flower initiation and arrested flower bud development in 'Pollyanne'. 125 Copper
deficiency depresses the activities of phenolase and IAA oxidase. The former effect allows
phenolic compounds to accumulate which can inhibit the action of IAA oxidase.' 24 It is
suggested that the cumulative effect of these responses might be to allow IAA to accumulate
within the plant and inhibit flower initiation (see below). 124

GROWTH REGULATORS

GAi 126..27 other GAS; i27 BA, 127 steroids, 128 and GA-like129 and sterol-like130 compounds
extracted from chrysanthemums committed to flowering, have all been shown to cause a
higher proportion of plants to form flower buds when the compounds are applied in LD.
Assessments based solely upon the numbers of plants with flower buds visible in a given
time are unsatisfactory, however, because flower buds are produced eventually on all chry-
santhemums grown in LD. Since some untreated plants had flower buds which were visible
by the end of many of the above experiments, it suggests that the initial populations were
not of a uniform physiological age.24-33 Although there is some evidence, therefore, that
GAs influence flower initiation in LD (and SD)' 3 ' and may represent the translocatable floral
stimulus that is produced in SD, 132 there are other reports that GAs do not promote flower
initiation. 130 - 133135
There is stronger evidence that GA3 can replace all or part of the chilling requirement
that some cvs have,"- 59 - 136 - 137 though again, some negative results have been obtained. l38 - 139
The concentration of endogenous GA in plant extracts rose after plants had been vernalized
and fell when they were transferred to low-light conditions which devernalized them, 137
while the application of chlorphonium chloride (Phosfon) to vernalized plants lowered their
GA content 137 and delayed flower initiation. 131 The latter effect could be reversed by the
simultaneous application of GA,. 131 Chlorphonium chloride, however, also affects the bio-
synthesis of sterols in tobacco, 140 and some sterols can influence flowering in
chrysanthemum. 12S - 130
GA, and other GAs clearly hasten anthesis when applied in SD to plants that have already
initiated flower buds, 134 - 141 l44 and promote floret differentiation when supplied to isolated
flower buds in LD. 145 Further evidence for the involvement of GA in flower development
 Volume II 251

comes from studies with growth retardants which inhibit stem extension and usually delay
flower development; both effects being reversed by the application of suitable GA. l 4 1 - 1 4 '- 1 4 4
In addition, extracts from apical buds of induced plants' 4 ' 1 and from young flower buds 147
contain high concentrations of GA-like compounds; their concentration declines later and
seems related to the relative growth rates of the developing buds. 147 The probable involvement
of GA in flower development emphasizes again the difficulty in assessing effects on flower
initiation solely in terms of whether flower buds are visible.
It is evidently easier to demonstrate an effect of growth regulators in terms of inhibiting
flowering and there are many reports that auxins alone,'"•" 5 - l48J4 '' or in combination with
GA,,' 2 can delay flower initiation in SD. They may also delay flower bud development, 5 - 44 l48
though this is not always observed. 1 4 I J 4 2 - I 4 > ' Extracts from plants in LD contain higher auxin
levels than those in SD, 44 - 1 "- 150 and so there is a reasonable body of evidence that the
inhibitor of flowering produced by leaves in LD"-' 5 ' is an auxin.
The most potent inhibitors of flower initiation are the unsaturated hydrocarbon gases,
ethylene and propylene. Continuous exposure to an atmosphere containing between 1 and
4 vpm ethylene inhibited flower initiation in SD,' 52 keeping plants vegetative throughout a
12-week period of treatment. 1 " Flower buds were formed when ethylene was supplied
intermittently, but their further development was retarded. 152 •'" Continuous exposure to
atmospheres containing 500 vpm propylene inhibited flower initiation both in SD and LD. 154
Thus propylene, and probably ethylene also, can block the photoperiodic and autonomous
flower-inducing processes.
Effects similar to these have been obtained by spraying plants with the ethylene-generating
compound ethephon at concentrations of from 100 to 2000 ppm in SD 155 - 156 and LD. 1 5 6 -' 5 7
The effects of ethephon persist for some time after treatment, perhaps because the plant's
tissues are stimulated to produce ethylene, but those of ethylene and propylene gases appear
to be dissipated as soon as the gases are removed from the atmosphere, 153 - 154 although the
tissues of treated plants do contain higher levels of auxins. 152 Ethylene may contribute to
the regulation of flowering under natural conditions, for treated plants take on a resetted
appearance152 l54 which is reminiscent of that of devernalized shoots from cold-requiring
cvs.

FLOWER ABNORMALITIES

Virus infections can cause flower abnormalities 1 and it is suspected that mycoplasma-like
organisms may cause the production of green flowers in some cvs by suppressing floret
appearance while encouraging bract formation over the whole surface of the receptacle. 15 "
The most common flower abnormality is the "crown bud", in which bud development is
severely retarded and the involucral bracts become grossly enlarged. 5 - 68J5y Crown buds are
produced on plants grown only in LD or transferred to LD after receiving a few SD (see
above).
Other abnormalities commonly occur when the period of floret formation is spent in
unfavorable conditions. Transfer to LD at this stage encourages the production of more ray
florets 7 " 9 - 16 - 44 - 68 and so enhances doubleness,"'" while SD enhance singleness. 8958 Similar
effects are observed if high temperatures (>30°C) are given during SD. 9 - 15 - 102 Transfer to
LD can also cause floret distortion68-161 and the production of long, tubular florets'60 which
are also seen following auxin treatment" or transfer to low temperature in some cvs. 162 In
addition to these abnormalities, bracts may appear on the receptacle in LD 5 • 6 - 16 - 44 • K)l - 163 and
at high temperatures, 102 and one or more secondary inflorescences may be formed within
the original flower bud. 5 - 16 - 44 - 161 -164 In some cvs these secondary inflorescences arise from
floret primordia.5 44
252 CRC Handbook of Flowering

REFERENCES

1. Cathey, H. M., Chr\santhemum morifolium (Ramat.) Hemsl., in The Induction of Flowering, Evans, L.
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29. Langton, F. A., Chimerical structure and carotenoid inheritance in Chrysanthemum morifolium (Ramat.).
Kuphytica, 29, 807—812. 1980.
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33. Langton, F. A. and Cockshull, K. E., Screening chrysanthemums for leaf number in long days, Annu.
Rep. Glasshouse Crops Res. Inst. 1978, 177—186, 1979.
34. Langton, F. A., Royce, H. M., and Cockshull, K. E., Leaf numbers of chrysanthemum cultivars grown
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35. Langton, F. A. and Dixon, T.,Annu. Rep. Glasshouse Crops Res. Inst. 1982, 104—127, 1982.
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37. de Jong, J., The difference between chrysanthemum plants raised from seeds or cuttings in their response
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translocation of the stimulus, J. Exp. Bot., 5, 389-^00, 1954.
39. Vince, D. and Mason, D. T., The effect of removing apical sections of the stem on the flowering behaviour
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500, 1981,
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Congr., Vol. 2, 1955, 1023—1039.
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45. Cockshull, K. E. and Hughes, A. P., unpublished data, 1970.
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4131, 484-^93, 1975.
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temporary chilling, J. Exp. Bat., 1, 329—343, 1950.
50. Mason, D. T., Chrysanthemum. I. A classification of some mid-season and later flowering varieties
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51. Schwabe, W. W., Factors controlling flowering in the chrysanthemum. V. Devernalization in relation to
high temperature and low light intensity treatments, J. Exp. Bot., 6, 435—450, 1955.
52. Schwabe, W. W., Effects of temperature, day length and light intensity in the control of flowering in the
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53. Mason, D. T. and Vince, D., The pattern of growth in chrysanthemum as a response to changing seasonal
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254 CRC Handbook of Flowering

61. Charles-Edwards, D. A., Cockshull, K. E., Horridge, J. S., and Thornley, J. H. M., A model of
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73. Cathey, H. M. and Borthwick, H. A., Phytochrome control of flowering of Chrysanthemum morifolium
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75. Ben-Jaacov, J. and Langhans, R. W., 'After lighting' of chrysanthemums. N.Y. State Flower Growers
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77. Borthwick, H. A. and Cathey, H. M., Role of phytochrome in control of flowering of chrysanthemum,
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78. Cathey, H. M. and Borthwick, H. A., Significance of dark reversion of phytochrome in flowering of
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79. Sachs, R. M., Kofranek, A. M., and Kubota, J., Radiant energy required for the night-break inhibition
of floral initiation is a function of daytime light input in chrysanthemum, HortScience, 15, 609—610. 1980.
80. Cockshull, K. E., Controlling bud formation. Grower, 93(8) (21.2.80), Suppl., 19—20. 1980.
81. Canham, A. E., The fluorescent tube as a source of night-break light, Exp. Hortic., 16. 53—68, 1966.
82. Kadman-Zahavi, A. and Yahel, H., Phytochrome effects in night-break illuminations on flowering of
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83. Accati-Garibaldi, E., Kofranek, A. M., and Sachs, R. M., Relative efficiency of fluorescent and
incandescent lamps in inhibiting flower induction in Chrysanthemum morifolium "Albatross", Ada Hortic.,
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84. Stuart, N. W., Controlling time of blooming of chrysanthemums by the use of lights, Proc. Am. Soc.
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85. Post, K., It's a short night you want, N. Y. State Flower Growers Bull., 99, 3—4, 1953.
86. Cathey, H. M,, Participation of phytochrome in regulating internode elongation of Chrysanthemum mor-
ifolium (Ramat.) Hemsl., J. Am. Soc. Hortic. Sci., 99, 17—23, 1974.
87. Cathey, H. M. and Campbell, L. E., Effectiveness of five vision-lighting sources on photo-regulation of
22 species of ornamental plants, J. Am. Soc. Hortic. Sci., 100, 65—71, 1975.
88. Cathey, H. M. and Borthwick, H. A., Photoreactions controlling flowering of Chrysanthemum morifolium
(Ramat. and Hemsl.) illuminated with fluorescent lamps. Plant Physio/., 45, 235—239, 1970.
89. Cockshull, K. E. and Prue, D. \.,Annu. Rep. Glasshouse Crops Res. Inst. 1979, 64—66, 1980.
90. Kadman-Zahavi, A. and Ephrat, E., Effects of red and far-red illumination at the end of short days and
their interaction with nightbreak illuminations on flowering of Chrysanthemum morifolium plants, Plant
Cel/Physiol., 14,409-^11, 1973.
 Volume II 255

91. Kofranek, A. M., Experiments continue with cyclic lighting for greenhouse mums, Flor. Rev.. 133(3434),
23—24. 1963.
92. Kofranek, A. M. and Carmeli, C., Cyclic lighting tor greenhouse chrysanthemums. Calif. State Flor.
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