A better way to construct the sunflower head
A better way to construct the sunflower head
HELMUT VOGEL
Technische Unicersitiit Mikehen, Lehrstuhl fiir
Physik,
D-8050 Freising- Weihemtephan, Federal Repiblic of Germany
ABSTBACI
INTRODUCTION
The flower heads (capitula) of most compositae consist of a great
number of individual flowers, whose arrangement has intrigued observers at
least from 1754 on [2]. The most conspicuous feature of this pattern is two
sets of arcs, one running clockwise, the other counterclockwise, and com-
posed of particularly tightly packed seeds. These arcs are often (wrongly)
described as logarithmic spirals [3-51. Their number in each set is invariably
a member of the Fibonacci sequence 1, 1,2,3,5,8,13,21,. . . ; e.g. 21 and 34
for a small sunflower capitulum, up to 89 and 144 or even 144 and 233 for
big ones.
The conspicuous arcs are not the basic structural elements, but merely
consequences of a single primary growth spiral on which, in the developing
capitulum, new individual flowers are added from the center (the
meristema), each issuing at an angle of 137.5” with respect to the preceding
one. This 137.5” angle occurs also in the arrangement of leaves or side
branches around the main stem of many plants, especially compositae,
where it leads to a near-periodicity in Fibonacci distances: The 5th branch
is nearly above the 0th; the 8th and 13th are even nearer to it. Sometimes
one can pursue this sequence up to the 55th branch, e.g. with the steppe and
dump plant mugwort (Artemisia). Thus, the growth program does not have
to change from the stem to the capitulum as far as angles are concerned,
but the longitudinal growth is relieved by a lateral one.
For convenience of description, I will count the flowers or seeds outward
from the center, i.e., contrary to their seniority. In polar coordinates r, cp,
flower number k is at q+ = k8 (6 = 137.5”) and at r, = r(rpk). What the lateral
growth program [i.e., the function r(q)] is and how it produces the flower or
seed pattern seems still to be controversial. Almost any r(q) explains the
existence of the arcs. Finer features are more discriminating, e.g. the facts
pointed out by Richards [6, 71 and Davis, Mathai [I]: The arcs do not run
all the way to the center, but are successively relieved by other systems of
arcs, fewer, but always Fibonaccian, in number; at the junctions of different
arc systems, irregular seed shapes arise. Davis and Mathai [1] proposed a
graphical construction that well describes these features, and a computa-
tional procedure that claims to do so. The former is based on the assump-
tion that the area provided for individual flowers or seeds is the same all
over the capitulum, the latter on the assumption that the primary spiral is
logarithmic (r = r,e”v) and because of its flatness (small a) approaches an
Archimedes spiral rmr,( 1 + acp). Confirmation is derived from the construc-
tion, not from the calculation, which I will show to contradict that construc-
tion and the facts.
I will provide a growth program that is biologically reasonable and
predicts in a quantitative and concise form the whole pattern of the
sunflower head down to the most minute detail.
WHY 137.5”?
The angle of 137.5”, common to stem and capitulum, can be considered
as the consequence of a very simple growth program. We look at the stem in
its axial direction and mark the side branches as points on a circle. Between
them, in general, there remain gaps of different sizes. The program then
prescribes:
(1) Each new branch (leaf, flower) bears the same spatial relationship to
the existing stem and accordingly issues at a fixed angle 6 with respect to
the preceding branch.
(2) Each new branch fits into the largest still-existing gap between older
branches, cutting a constant fraction off that gap.
Pn+l W”
zn+,=-=
%+I q,z-1 ’
(1)
which is generally true, starting from z,,= f, if and only if pn and qn are
consecutive Fibonacci numbers of even order:
r(cp)=a*. (4
FIG. 1 The primary cyclotron spiral connecting consecutive seeds that have a
Fibonacci angular distance of 137.5”.
or seeds. From (2) for the distance dk, between the centers of seeds k and I
we have, according to the cosine theorem,
dk:
aZ=k+l-2flcos(k-I)S. (3)
x=k-lxs: 2’
&27T (4)
g ET2 .
x=F,, m= Fn_2.
This explains the fact that in an arc, neighbors have always numbers
differing by a Fibonacci x. Of course, (4) becomes an exact equality only in
the limit, when F,_,/ F, attains g2. The difference x&m-277 is
1.073
Gg2- F,-z= 2,rFn_2. (5)
In the interesting region, one can thus expand the cosine around rna2r:
d2
‘ak+l-2m(l-$$),
a2
dk:
-= x2 p2k x3 p2 1.073
4k+-+s-p P-- - =2.809 (6)
a2 x2 g2
TABLE 1
One arc extends over an angular distance of about 1, i.e., 60”. Its inclination
with respect to the radius, arctan(rdq/dr), increases from 20” at the inner
end to 67” at the outer one. Thus, the angle between arc and radius changes,
which already suffices to show that the arcs are not logarithmic spirals, for
which this angle would be constant. (Compare the flight of nocturnal insects
around a street lamp: Constancy of angle between flight and light direction,
which normally leads to a straight course, here produces logarithmic
spirals.)
186 HELMUT VOGEL
All these details are confirmed equally well from actual capitula and
from constructions according to the cyclotron program (cf. Fig. 2). Davis
and Mathai’s [l] graphical (not their computational) procedure also fits
reasonably well, apart from inaccuracies which may be due to drawing. The
quasi-identity of the two constructions is not surprising, since Davis and
Mathai also use the Fibonacci angular condition and approximate the
equal-area condition by pushing circles or coins of equal size along their
Fibonacci radii until the new one touches at least one of the previous ones.
Here,
d2
-$ = k2”+ 12”-2k”l”cos(k- /)a,
e -bx=l.---_,
P’ P2
bx’ 2x2
BETTER WAY TO CONSTRUCT THE SUNFLOWER HEAD 187
FIG. 3 A capitulum of 959 seeds, based on the Archimedes growth program instead
of a cyclotron one.
dm=aebk(b-m)mabe”.
x,,, is independent of k, i.e., there is only one type of arc in either sense,
which runs all the way from the center to the periphery. The distance d,,,
heavily depends on k. Here and only here, the arcs are logarithmic.
Finally, we consider a program that ensures termination of lateral
growth, with still constant frequency of appearance of individual flowers:
qk = k& Iji=a(l-emck),
where k this time has to count in the order of seniority, i.e., small k at the
188 HELMUT VOGEL
d,~a~ee-2ckc2x2+~2x~2(1-e-ck)2 ,
For ck<<l, one obtains of course the Archimedes-spiral results, and for
ck>>l
Seeds with ck>l (i.e., at less than 30% of the final radius a of the
capitulum) would become smaller and smaller, and the arc structure would
become indistinct, since v- runs through the Fibonacci sequence
very quickly.
DISCUSSION
From a teleological point of view, the problem of the sunflower head can
be considered settled. Two simple principles-most uniform angular dis-
tribution of branches and flowers (hence the Fibonacci angle), and most
uniform area distribution of flowers and seeds (hence the cyclotron primary
spiral)-explain the pattern down to the most minute apparent irregulari-
ties. The Fibonacci law permits the angular growth program to be the same
for stem and flower head; the cyclotron law prefigures the transition from a
growing flower head to the final state of this head. What has still to be
provided is a causal mechanism for both growth programs.
Agronomists use the arc structure as an aid in counting the seeds on a
capitulum, e.g. in resolving the crop yield into its components (number of
plants/acre, number of flowers/capitulum, percentage of flowers that bear
seeds, weight of seed, oil content), and in studying the he&abilities of these
components [8]. Irregularities in the arc structure, especially the fact that
arcs do not run all the way through, slightly falsify simple counts according
to the schema (number of peripheral arcs)x(number of seeds per arc).
More important, it is conceivable that regions of abortive flowers (produc-
ing no seeds or empty seeds), which one observes on actual capitula and
which can reduce the yield to 60% or less, are related to the regions where
one arc system is relieved by another [9]. The nature of this relationship,
and whether it is simply due to deformed flowers in the transition regions
being avoided by bees, are under study.
REFERENCES