Stems and cones

A disc is a cylinder

Vogel's seed head makes the rise fall as one over radius squared, so a disc is not one lattice but a family of them. Feed that into the cylinder's ladder and it predicts where a sunflower's spiral counts change — with nothing fitted, and against a counter that never sees either model.

Worth reading first: The Fibonacci ladder.

The strongest thing a figure on this site can do is arrive at a number twice, by arithmetic that shares nothing. This essay is the clearest instance of it the site has.

On one side is a lattice on a cylinder: a periodic strip, two parameters, and a ladder of parastichy pairs as one of them falls. On the other is Vogel’s disc: the nn-th primordium at angle nδn\delta and radius cnc\sqrt n, which is where the seed-head figures come from and which has no rise in it anywhere.

The two share the divergence angle and nothing else. No constant is fitted between them. And the cylinder’s ladder predicts, to a few per cent, the radii at which a blind counter run on the disc changes its answer.

Where the disc's counts change, predicted from a cylinder. The dashed lines are the transition radii the cylinder's ladder gives through h = c²/4πr², with nothing fitted. The dots are what the blind counter returns from the disc: 15 of 16 bands agree, and the ones that do not sit on a transition.
Fig. 1 The dashed lines are transition radii computed from the cylinder’s ladder. The dots are what the disc’s own counter returns, band by band, having seen neither the cylinder nor the divergence angle. Fifteen of sixteen bands agree.

The rise of a seed head

The derivation is three lines and it is the whole essay.

In Vogel’s model primordium ii sits at radius r=cir = c\sqrt i. Differentiate: between one primordium and the next, the radius advances by

drdi=c2i=c22r\frac{dr}{di} = \frac{c}{2\sqrt i} = \frac{c^2}{2r}

The local circumference is 2πr2\pi r. The rise — the vertical advance per element, measured in circumferences, which is the cylinder’s second parameter — is therefore

h(r)=12πrc22r=c24πr2h(r) = \frac{1}{2\pi r} \cdot \frac{c^2}{2r} = \frac{c^2}{4\pi r^2}

That is it. A disc has a rise, it is not constant, and it falls as the square of the radius.

What that means about seed heads

Three consequences follow immediately, and together they replace an observation with an explanation.

A disc is not one lattice. Each annulus is a cylindrical lattice at its own rise, slightly smaller than the one inside it. The head is a continuous family, indexed by radius.

The counts must change. Which is what the founding essays found by counting, without being able to say why. A falling rise climbs the ladder, so the parastichy pair at the rim is higher than the pair near the centre. This is not a special property of seed heads; it is what any cylinder does when its rise falls, and the disc is simply a geometry in which the rise falls very fast.

The changes are at computable radii. Inverting the relation gives

r=c2πhr = \frac{c}{2\sqrt{\pi h}}

so each transition rise on the cylinder’s ladder maps to a transition radius on the disc, with cc the only quantity involved and cc being the disc’s own scale rather than a fitted parameter.

The foundation essay The counts change with radius established the phenomenon by counting. It did not have this, and could only say that the transitions happen. Now they can be placed.

The prediction, and the test

The cylinder’s transitions at the golden divergence sit at rises of 0.1246, 0.0476, 0.0182, 0.00695, 0.00266 and so on downwards, each 0.382 of the one before.

Convert with r=c/2πhr = c/2\sqrt{\pi h}, at c=1c = 1: transition radii of 0.80, 1.29, 2.09, 3.38, 5.48, and continuing upward by a factor of φ\varphi each time — 8.87, 14.4, 23.2, 37.6.

Now the test. A disc of 3,000 points at the same divergence is cut into sixteen radial bands, and each is handed to the disc’s own parastichy counter — the one that receives coordinates and a radius window and nothing else. Its answers are compared with the pair the cylinder says should hold at the rise corresponding to that band’s mid-radius.

Fifteen bands of sixteen agree. The pairs run 13/21, 21/34, 34/55, 55/89, 89/144 as the radius grows, and the predicted and measured sequences match band for band.

How much of that agreement survives on a smaller head is a separate question, and it costs nothing to ask. The prediction is the same curve whatever the head holds, because it is computed from the divergence and the radius and knows nothing about how many points there are; what the size decides is how far out a blind counter can still find chains, and therefore how many of the predicted transitions the test gets to reach at all. So the same test is made at six head sizes, from three thousand points down to nine hundred, and the six differ only in how many rungs of the ladder they are able to put a dot on.

Where the disc's counts change, predicted from a cylinder. The dashed lines are the transition radii the cylinder's ladder gives through h = c²/4πr², with nothing fitted. The dots are what the blind counter returns from the disc: 15 of 16 bands agree, and the ones that do not sit on a transition.
Fig. 2 The same test on a head of two thousand four hundred points. The dashed transitions are where they were, because the prediction does not know the head’s size; what a smaller head changes is how far out the counter still returns a pair.

The one that disagrees

The disagreement is at a mid-radius of 37.1, where the counter returns 89/144 and the cylinder says 55/89.

Where the disc's counts change, predicted from a cylinder. The dashed lines are the transition radii the cylinder's ladder gives through h = c²/4πr², with nothing fitted. The dots are what the blind counter returns from the disc: 14 of 16 bands agree, and the ones that do not sit on a transition.
Fig. 3 Two thousand points. The outer bands are the ones a disagreement can happen in, because they are the ones where consecutive transitions are close together in radius, and they are also the first bands a smaller head loses.

The predicted transition between those two pairs is at 37.6. The band that disagrees straddles it.

This is the least surprising possible failure and it is worth not smoothing over, because a prediction that agreed in sixteen bands of sixteen would be less informative. Right at a transition three offsets have nearly equal hops; the counter’s answer there is decided by where the band edges fell and how many points landed inside them; and the model’s answer is decided by whether the mid-radius is a hair above or below a threshold. Both are correct and they are answering a question that has no sharp answer at that radius.

Any measurement that never disagreed with its prediction anywhere would be a measurement with no resolution.

What c is, and why it is not a knob

The conversion carries one symbol from the disc into the cylinder’s world, and it is worth being clear that it is not a free parameter dressed up as a constant.

In Vogel’s model, cc is the only scale there is: the nn-th primordium sits at cnc\sqrt n, so cc fixes how much area each element gets. Since the area of the disc out to primordium nn is πc2n\pi c^2 n, the area per element is exactly πc2\pi c^2 — which makes cc a statement about seed size relative to head size and nothing else.

It is therefore measurable on a head without reference to any of this. Count the elements inside a radius, divide the area by the count, and c=A/πnc = \sqrt{A/\pi n}. The disc recovery in the round-trip essay already does exactly that, estimating cc from the density in the counted band rather than taking it from the model, precisely so that the recovery is not handed its own answer.

So the prediction here uses a quantity that comes out of the point positions. If it had to be fitted — if the transition radii could be moved to match the measurements by adjusting cc — the agreement would mean very little. It cannot: change cc and every predicted radius moves together, by the same factor, so the spacing of the transitions is untouched and only their absolute position shifts. The spacing is a factor of φ\varphi per rung and it is not adjustable at all.

Running it backwards

If a rise predicts a pair, a pair constrains a rise, and that turns the relation into a measuring instrument.

Suppose one has a photograph of a head, counts 34 and 55 at a stated fraction of the radius, and wants the head’s scale. The pair pins the rise to the interval between two transitions — for 34 and 55 that is 0.000149 to 0.000390 — and the radius at which it was counted then gives cc to within a factor of 0.000390/0.000149=1.6\sqrt{0.000390/0.000149} = 1.6.

Where the disc's counts change, predicted from a cylinder. The dashed lines are the transition radii the cylinder's ladder gives through h = c²/4πr², with nothing fitted. The dots are what the blind counter returns from the disc: 13 of 15 bands agree, and the ones that do not sit on a transition.
Fig. 4 Twelve hundred points, which is the regime the photograph argument is actually in. The scale recovered from one counted pair is only as good as the band it was counted in, and this is how many bands a head that size offers.

That is not a precise measurement, and the imprecision is honest: a pair of counts is a coarse observation, and a range of rises produces it. But it is an independent one. The usual way to get cc from a photograph is to count elements and measure area, which requires resolving every element; this way requires only tracing two families of spirals, which is what a person can do on a bad photograph of a distant flower.

Two routes to a scale, one of them cheap and coarse and the other expensive and precise, disagreeing by less than the coarse one’s stated range. That is the same arrangement as everything else in this field, run in the direction that is actually useful for measuring things.

Why this is not circular

It is worth being precise about what would make this trivial, because the arrangement invites suspicion.

It would be circular if the disc counter used the cylinder’s arithmetic. It does not: the disc counter measures hop lengths between points in an annulus, normalises by the local spacing, and takes local minima of the resulting curve. There is no wrap, no rise, and no lattice vector anywhere in it.

It would be circular if cc were fitted. It is not: cc is the disc’s own scale factor, fixed when the point set is built, and the conversion uses it as it stands.

It would be circular if the two models shared a derivation. They do not. The cylinder’s ladder comes from comparing jδ2+(jh)2\sqrt{\langle j\delta\rangle^2 + (jh)^2} across jj. The disc’s counts come from medians of Euclidean distances between points at cic\sqrt i and ci+mc\sqrt{i+m}. The n\sqrt n is in one and not the other; the wrap is in the other and not the one.

What they share is the divergence angle, which both are given, and which by itself determines neither answer.

What the agreement buys

Two things, one about the models and one about the subject.

About the models: the cylinder is not a separate topic that happens to sit next to the disc in this collection. It is the general case, and the disc is a particular trajectory through it — the trajectory hr2h \propto r^{-2}. A cone is a slower trajectory. A stem is a stationary point. Everything the site has said about seed heads is a statement about one path through the two-parameter plane, which is why it kept needing qualifications.

About the subject: this is the cleanest available answer to why sunflower counts are Fibonacci numbers at every radius rather than only at one. The head does not settle on 34 and 55; it passes through 13/21, 21/34, 34/55 and 89/144 on the way out, and it does so because the rise falls monotonically and the ladder has rungs at the convergent denominators. The famous pair is whichever rung the counting band happened to land on.

How fast a head climbs

The geometric spacing of the transitions has a consequence specific to discs, and it is a number.

Consecutive transitions are a factor of φ2\varphi^2 apart in rise, and hr2h \propto r^{-2}, so they are a factor of φ1.618\varphi \approx 1.618 apart in radius. Every time a head’s radius grows by 62%, its counts advance one rung.

That is a slow climb and it explains something about how the pattern reads. The outer half of a large head spans a factor of two in radius, which is one rung and a bit — so most of the visible surface of a sunflower shows a single pair, with the changes squeezed into the crowded middle where nobody counts. Hence the impression that a head has a pair, and hence the surprise when someone counts carefully at two radii.

Where the disc's counts change, predicted from a cylinder. The dashed lines are the transition radii the cylinder's ladder gives through h = c²/4πr², with nothing fitted. The dots are what the blind counter returns from the disc: 13 of 15 bands agree, and the ones that do not sit on a transition.
Fig. 5 Sixteen hundred points. The number of dashed lines the dots reach across is the number of transitions a head this size is able to show at all, which is the same slow climb counted rather than described.

It also puts a bound on how many transitions a head can show. A countable region spanning a factor of four in radius holds about three, which is what the three-regime figure finds directly.

The cone in between

A cone is the intermediate case and the relation makes it precise.

On a cone the circumference grows linearly with distance from the apex, so with fixed internode spacing the rise falls as 1/z1/z rather than as 1/r21/r^2 — one power rather than two. Transitions are therefore a factor of φ22.618\varphi^2 \approx 2.618 apart along the axis rather than a factor of φ\varphi, and a cone of ordinary proportions shows fewer of them than a disc of comparable extent.

Which matches what one sees. A pineapple holds 8 and 13 over most of its length; a sunflower changes pairs two or three times across its face; and a stem holds one pair for ever. The two objects are the same lattice family sampled along two different paths, and the difference in how often they change is the difference between one power and two.

Every transition as the rise falls. The pair climbs 1/2 → 2/3 → 3/5 → 5/8 → 8/13 → 13/21. Consecutive transitions are 0.382, 0.382, 0.383, 0.382 of the previous rise — 1/φ² is 0.3820.
Fig. 6 The ladder both objects are moving along. A disc travels it at a factor of φ in radius per rung; a cone at a factor of φ² along its axis; a stem does not travel it at all.

What a real head does that neither model does

The agreement is between two models. Real heads depart from both, in ways that are known and that the prediction should be read against.

The centre is wrong. Vogel’s n\sqrt n gives infinite density at r=0r = 0, and a real head has a finite disc of tiny undifferentiated primordia there. Every count taken inside about a tenth of the radius is a count of something the model does not describe, which is why the bands in the test start at 0.10 of the radius rather than at zero.

The rim is wrong. Real seeds are packed against a boundary, are often flattened against it, and the outermost ring is frequently incomplete. The last band in the test is the one most likely to be measuring an artefact of the model’s edge rather than a lattice.

Seeds are not points. They have size, and they compete for space, so a real head is closer to a packing than to a lattice of centres. The effect is small where the pattern is regular and is not small where it is not.

None of these disturbs the prediction, and there is a reason: the prediction is about which offsets give the shortest hops, which is an ordering rather than a distance, and an ordering survives a good deal of distortion before it changes. That robustness is why parastichy counts are a usable measurement on real material at all, and it is the same reason the counts are hard to make precise near a transition, where the ordering is nearly a tie.

Where the disc's counts change, predicted from a cylinder. The dashed lines are the transition radii the cylinder's ladder gives through h = c²/4πr², with nothing fitted. The dots are what the blind counter returns from the disc: 12 of 14 bands agree, and the ones that do not sit on a transition.
Fig. 7 Nine hundred points, the smallest head tested here. The prediction is unchanged and the agreement is asserted on it exactly as at three thousand; what a head this small costs is the outer bands, not the ordering.

What this does not settle

The prediction says where the counts change. It does not say why the divergence is 137.5°, and nothing in this essay bears on that question — the angle is an input to both models.

It also does not say anything about mechanism, and what a mechanism would have to show is a longer list than either model touches. Both models describe form. A plant that produces a pattern matching Vogel’s disc is not thereby shown to compute cnc\sqrt n, and this site is careful about that everywhere because the same patterns come out of magnetised droplets with no biology in them.

What has been established is narrower and more useful than either: that two descriptions of phyllotactic form, built from different arithmetic, are descriptions of the same object. When a model’s prediction lands on a measurement made by machinery that never saw it, the number is not an artefact of either — and on a subject where almost every quoted figure is an artefact of how it was quoted, that is worth having.

One counter, three surfaces. A cylinder's counter returns 3/5 in every band and never changes. A cone's transitions are spaced by 2.658 against φ² = 2.618; a filling disc's by 1.645 against φ = 1.618. Equal spacing on a log axis is what a geometric ladder looks like.
Fig. 8 The cone this essay predicts in its last section, now computed. One counter on three surfaces: a cylinder that never changes, a cone whose transitions are a factor of φ² apart, and a filling disc whose are a factor of φ.

One more thing the derivation gives away

The relation hr2h \propto r^{-2} has a consequence that is easy to miss and that reframes a piece of the founding essays.

The rise falls without limit as the radius grows. There is no smallest rise, so there is no highest rung, so a sufficiently large head has arbitrarily high parastichy numbers — 233 and 377, and beyond, with no change in the model and no new physics. The Fibonacci numbers people quote are the ones that fit on a flower, not the ones the model favours.

The other direction is more interesting. As r0r \to 0 the rise grows without bound, and above a rise of about 0.125 the dominant pair is 1 and 2 — the lowest rung there is. So the very middle of a head is, in the model’s terms, a distichous pattern: two families, one of one spiral and one of two, which is to say almost no pattern at all.

That is not visible in a real flower, because the middle of a real flower is the undifferentiated disc the model does not describe. But it is a statement the model makes, it follows from arithmetic already on the page, and it is the sort of thing worth recording when a derivation turns out to say more than it was asked.

It also has a check attached, which is what stops it being a curiosity. If the disc really is a cylinder with hr2h \propto r^{-2}, then the rises at which its counts change are the cylinder’s own transition rises — a discrete set with a closed form — mapped through that relation into radii. So the model predicts where on a head the counts should change, not merely that they should, and the prediction is a list of numbers rather than a trend. Run a blind counter outward across a head and compare: agreement in fifteen bands of sixteen, with the one disagreement sitting on a transition.

What links here

Computed from the collection, not written here: the essays that point at this one.

Reads more easily once this is understood

Essays that name this one as worth reading first.

Shares its objects with

Essays that name at least two of the same things, and that neither author linked.

Named objects

A flat tag is an object no other essay names yet.

Cross validationCylinderDivergence angleLadderParastichyParastichy pairPredictionRiseTransition radiusTransitionsVogel's model