Phyllotaxis
the spirals are counted, not admired
Concept

Parastichy — where it appears

One of the families of spirals running through a phyllotactic pattern, named by how many places apart its consecutive elements are. The families worth counting are the two whose steps are shortest, and which two those are changes with radius on a single head.

Named by 35 essays across 6 fields — each of them below, with the objects they name alongside it.

A stem unrolled: 28 nodes at 137.51° with a rise of 0.090 circumferences. The counter is shown these coordinates and the circumference, and finds 2 parastichies one way and 3 the other. The faint strips left and right are the same stem: a family leaving one edge re-enters at the other.

A stem is a cylinder

The sunflower is the photograph, and it is the hard case. Nearly all real phyllotaxis happens on a stem, where the geometry is a lattice on a cylinder with two parameters — and where the spiral counts, which on a disc change with radius, are the same the whole way up.

cylinder · Stem lattice A head of 200 primordia at a divergence of 137.51°. Nothing is placed by hand: the nth point sits at n·137.51° and radius √n. The closest any two points come is 1.60 of the mean spacing.

A head is a set of points

The nth primordium at n times an angle, and a radius of root n. Two lines of arithmetic produce a sunflower head, which is either remarkable or suspicious depending on how carefully the claim is stated — and stating it carefully is most of the work.

lattices · Lattice The two spiral families a counter finds between 0.68 and 0.92 of the radius. 34 spirals one way and 55 the other, found from the point positions alone — the counter is never told the divergence angle.

Counting the spirals

Almost every claim about phyllotaxis is a claim about how many spirals run through a pattern, and the count is almost never done. It can be done from the points alone, by a count that is never told what angle built them — and then a count of 34 is evidence rather than a restatement.

lattices · Counting The same counter, on a stem and on a disc. The stem returns 2 and 3 in all three bands. The disc returns 21/34, 34/55, 55/89 — three answers to one question, which is why a published count needs to say where it was taken.

Counting up the stem

The same counting machinery, pointed at a stem instead of a seed head, returns one answer three times where the head returned three answers. That contrast is a measurement rather than a preference, and it is the one the whole cylindrical argument rests on.

cylinder · Stem count Six stems built, forgotten and recovered. Each row is a lattice built from a divergence and a rise, counted by machinery shown only the coordinates, and reconstructed from the counts and the two hop lengths. The worst error in the recovered angle is 3.0e-13°.

Two numbers out of the points

A seed head's divergence angle can be recovered from its spiral counts only to within an interval, because a range of angles gives the same counts. On a stem the counts come with lengths attached, two measurements pin two unknowns, and the lattice comes back to the last digit it was built with.

cylinder · Cylinder recovery A round trip on four heads of 400 primordia: the divergence angle recovered from each. The counter is shown the points and nothing else. The worst recovery across the four is 0.035°.

Recovering the angle from the counts

Build a head at a stated divergence angle, forget the angle, and get it back from the spiral counts alone. Four angles, worst error twelve thousandths of a degree — and the only thing that crossed between the two halves was a list of coordinates.

lattices · Recovery 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.

The Fibonacci ladder

Lower the rise on a cylinder and the parastichy pair climbs — 1 and 2, then 2 and 3, then 3 and 5 — each rung the sum of the two before it. Nothing in the arithmetic mentions Fibonacci, the transitions sit at computable rises, and consecutive ones stand in the ratio 1/φ².

cylinder · Ladder Which patterns grow on a ring of circumference 0.80. Modes 2 to 8 have positive growth rates and mode 4 is fastest. Integrating the full equations from a disordered start gives 4 peaks.

What a mechanism would have to show

Every model of phyllotaxis comes with the caveat that reproducing a pattern is not explaining it. That is easy to repeat and hard to make precise. Here it is made precise — a list of what an account of phyllotaxis would have to establish, with each item marked according to whether the models drawn in these essays establish it.

mechanism · Mechanism claims 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.

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.

cylinder · Rise and radius Tracing one family: 4 chains. Every node is joined to the node one repeated displacement away, and the chains that result are drawn separately. There are 4 of them, which is the parastichy number of this family. No index of arrival was used anywhere, which is what lets the same count be made on a pattern where 2 primordia appear at once.

Counting without an index

A person counting spirals on a cone puts a finger on one scale, follows a family round, and counts how many distinct chains there are. That needs no order of arrival — and building it turns out to be strictly more general than the counter that reads the order of arrival, and to find a bug in the counting of a bijugate stem that nothing had caught.

lattices · Chain counting A cone unrolled: 105 nodes at 137.51°, sector 126°. The counter is shown these coordinates and the sector angle and nothing else. Near the apex it finds 3 and 5; near the base, 8 and 13. The rise falls as one over the distance from the apex, so the pair has to climb.

A cone has a rise that falls

A stem holds one parastichy pair for ever and a seed head changes its pair with radius. A cone does both — it is a cylinder whose rise falls as one over the distance from the apex, and the same blind counter that finds one answer up a stem finds four up a cone.

cylinder · Cone A cell's neighbours are its spiral families. Left: part of a 900-point golden, 137.508° head, with every contact between two cells drawn and coloured by the difference between the two nodes' placement indices. Right: the share each difference takes, across all 1903 contacts between interior cells. They are the parastichy numbers — 34, 55, 21, 89, 13, 8 — and the pair a person would count is 34 and 55. The six sides Euler forces are shared out among four of them, 5.72 edges per cell.

The six are the spirals

Label every contact between two cells in a seed head with the difference between the two nodes' placement indices. The labels are the parastichy numbers — 34, 55, 21, 89 — and the six sides Euler forces turn out to be about two from one family, one and a half from the next, and one each from two more.

tissue · Contact network The angles against the positions, rise by rise. three rises, five seeded stems each. A filled mark is a run whose angle readout returned the pair the position counter finds in the same stem; an open mark is a refusal. At 0.032 the counter says 3/5 and the angles agree on 0 of 5, refusing 5. At 0.013 the counter says 5/8 and the angles agree on 5 of 5. At 0.005 the counter says 8/13 and the angles agree on 5 of 5. The two instruments share no code path: one is given a list of angles, the other a list of coordinates.

A counter that sees no positions

This site has counted spirals two ways, and both were handed coordinates. A third counter is handed a list of angles and nothing else. It returns one number instead of two, it refuses more often, and where it refuses it would have been wrong every time.

lattices · Counting Every family but two is the sum of two others. Four heads and the contact families each actually has. An arc arrives at every family that is the sum of two smaller ones; the two with no arc arriving are the generators, and on every head they are the two smallest. whorled, 144°: 2, 3, 5 — golden, 137.508°: 8, 13, 21, 34, 55, 89 — Lucas, 99.502°: 11, 18, 29, 47, 76 — rational, 137.5°: 8, 13, 21, 34, 55, 89 — 137.0°: 8, 13, 21, 29, 50, 71, 92, 113. So once a pair is counted the rest is arithmetic, and a third counted family is a prediction rather than a second measurement.

Every family but two is a sum

A seed head has six spiral families and everybody reports two. That looks like a convention hiding information and it is the opposite — every family but the two smallest is the sum of two others, so a third count is a prediction rather than a measurement, and a check that catches a wrong pair.

lattices · Counting The exponent sets the spacing, and only the exponent. The curve is φ^(2/p), drawn from the ladder's ratio of 1/φ² and nothing else. The dots are measured: each surface built, a blind counter walked up its axis, the places its answer changed recorded. A cone that elongates and a paraboloid that fills sit on the same point at p = 1 — so the spacing identifies neither the shape nor the way material arrives, only their product.

The shape and the law

A cone's transitions are a factor of φ² apart and a disc's a factor of φ, and the temptation is to read the ratio as the shape. It is not. Five surfaces built and counted show that the ratio measures one exponent, and that the exponent is the shape multiplied by the way material arrives.

cylinder · Shape exponent Three organs, and the exponent each has at each place. Left: the meridian of a cone, an ogive of arc radius 12, and a spherical cap of radius 40, each scaled to its own length. Right: d log r / d log s along it. The cone sits at 1 the whole way; the ogive starts near 2 at its blunt tip and falls to 0.23 by the end of the 100 per cent shown; the cap starts at 1 and falls slowly.

An organ has no single exponent

The earlier work measured that a surface whose circumference grows as a power of arc length puts its transitions a fixed factor apart, and checked it on five surfaces. Every one of them had a single exponent, and no organ does — a fir cone is an ogive, whose exponent runs from 2 at the tip to nearly 0 at the shoulder.

cylinder · Varying exponent One wrecked stem, lag by lag — golden, rise 0.005, organ 6 back. How much each lattice hop moves from organ to organ in a stem that never repaired after a single removal, over its last 120 organs. The lag-one hop is the divergence itself and it swings by 55 degrees. The lag-4 hop swings by 0.12 degrees and sits 0.01 degrees from where the undisturbed stem put it, so that one family of the original lattice is still standing organ by organ. Its multiples inherit the same steadiness and nothing else comes within a factor of twenty. The block of angles this stem repeats has a period of 4, which is the surviving lag and not a coincidence.

A period that is not a count

Eighteen wrecked stems settle into a block whose period is one of their own spiral counts, and one settles into a block of four on a lattice counted 8 and 13. The odd one is not noise. It is the case that shows what the rule is actually conserving, and it is the reason this thread is about lattice steps rather than about spirals.

lattices · Counting The measurement is limited by the protractor, not by the plant. The peak falls as the reading error grows, and it falls by an arithmetic factor with nothing fitted: a position error enters two consecutive divergences with opposite signs, adding variance at every lag while the pattern's signal sits at one. At a quarter of a degree the readout is right on all 5 runs; at half a degree on 2; at a degree on 1. Below the dashed floor the peak is the largest of thirty noisy numbers rather than a measurement.

What the protractor has to be

The readout that names the parastichy number costs sixty internodes, which is cheap. It also needs every organ's position measured to better than a quarter of a degree, which is not — and the requirement follows from arithmetic rather than from care, so no amount of averaging relaxes it.

wrong · Sequence The statistic everybody reports is the one that cannot vary. Six arrangements of 900 points, from a whorled lattice to a set with no rule in it. The mean number of sides per cell is 5.97–6.04 on all six, because Euler's formula forces it. The mean squared departure from six runs from 0.023 to 1.83 — a factor of 79 — and the most hexagonal tissue in the set is the whorled one, at a rational angle.

What a summary throws away

Four statistics this collection has relied on turn out to be incapable of varying with the thing they describe — one is invariant to shuffling, one is fixed by a theorem, one is a parameter that stopped mattering, one is a fitted number selected into being wrong. In each case the second statistic was free and nobody had taken it.

wrong · Second statistic The order of the angles carries the count. Three stems, each held at a fixed rise so the pattern sits on one rung of the ladder. At a rise of 0.032 the positions count 3 and 5 spirals and the angles peak at 3; At a rise of 0.013 the positions count 5 and 8 spirals and the angles peak at 5; At a rise of 0.005 the positions count 8 and 13 spirals and the angles peak at 8. Each panel marks the peak and its multiples; the pale strip is what an uncorrelated sequence of this length gives.

The order carries the count

Take the divergence angles off a stem, throw away every coordinate, and autocorrelate what is left. The result is periodic at the smaller parastichy number — peaks at it and at every multiple of it. A list of angles, with no picture and no position in it, carries the spiral count.

cylinder · Sequence Two combs, at a rise of 0.005. The autocorrelation of 760 divergence angles from one stem held at a rise of 0.005. The filled teeth are the lags at multiples of 8; the open teeth are the second comb, at the same spacing offset by 5. Reading the spacing off the first and the offset off the second gives the pair 8 and 13, which is what the position counter reports for the same stem — from angles alone, with no coordinate anywhere in the calculation.

The second comb

The autocorrelation of a divergence sequence has peaks at the smaller parastichy number and at every multiple of it. It also has a second set of peaks, at the same spacing, offset by the difference of the pair — so a list of angles with no coordinate in it returns both numbers rather than one.

cylinder · Sequence 264 exceptions in a 2400-organ head, and eight circles. A golden, 137.508° head of 2400 organs, tessellated inside 86% of its radius. Every interior cell is drawn, and the 135 five-sided cells and 129 seven-sided ones are marked apart from the 1367 hexagons, and the pale circles are radii computed from the divergence angle through the lattice's third-shortest vector — nothing is fitted. Every defect sits within 0.64 of a cell spacing of one of those eight circles, and between them there is not one exception in hundreds of cells.

The defects lie on rings

The cells in a seed head that are not hexagons are not scattered through it. Every one of 264 sits within a cell of a radius computed from the divergence angle alone, the radii are a factor of φ apart, and between two of them lie 422 consecutive cells without a single exception.

tissue · Topological charge Two combs, at a rise of 0.013. The autocorrelation of 760 divergence angles from one stem held at a rise of 0.013. The filled teeth are the lags at multiples of 5; the open teeth are the second comb, at the same spacing offset by 3. Reading the spacing off the first and the offset off the second gives the pair 5 and 8, which is what the position counter reports for the same stem — from angles alone, with no coordinate anywhere in the calculation.

A harmonic is a step taken twice

The spectrum contains the larger parastichy number, their sum, and echoes of the smaller one, and no ranking of peak heights separates them. What separates them is arithmetic: a harmonic is a multiple of the spacing and a family is not, and the two kinds sit in different residue classes.

cylinder · Sequence Four ways to count the rule's neighbourhood, and one ordering. How many organs the placement rule is looking at, as its falloff exponent is swept from 1.5 to 6, by three different definitions. Counting the organs that carry half the profile gives 33, 12, 3, 2, 1. Counting the organs that carry nine tenths gives 182, 120, 30, 8, 3. Counting the organs that carry the weighted mean lag gives 68, 45, 21, 14, 11. They disagree about the size of the neighbourhood by a factor of 10 — the widest moves by 61 times across the sweep and the narrowest by 6.1 — and every one of them falls as the exponent rises. So no choice among them changes which rule is the deeper, and no redefinition can turn a result about the depth around.

The nearest organ is not the nearest neighbour

Rank the terms of the sum the rule minimises and read off which organs the biggest ones belong to. At every falloff exponent from 1.5 to 6 the answer is the same five: lags 13, 8, 5, 21 and 26. The organ placed immediately before is not among them, and counting the neighbourhood in organs was the wrong unit.

mechanism · Noise colour

Two rankings, one list

An essay in this collection claimed that the four shortest index hops on a seed head and the four largest shares of its cell walls are the same four numbers in the same order, and called the correspondence exact. Measured again from the same points, the two lists hold the same four families and order them differently, and they order them differently in five of the six bands the head can be read in.

tissue · Neighbour definition

Removing a neighbour costs least

Take away an organ that is a direct chain-neighbour of the growing tip and the next organ moves by under thirty-one degrees. Take away anything else inside the front and it moves by at least sixty-three. Thirty cuts, two groups, a factor of two between them and nothing in the gap.

mechanism · Ablation

One level and two exceptions

Inside a wrecked stem's period most residue classes sit at one level and a couple do not. On seventeen of the thirty cuts the exceptions are exactly two, equal and opposite to within five per cent — and on all seventeen they are neighbouring residues, which was not looked for.

mechanism · Damage shape

A twist is a divergence

Recovering a head's divergence angle from its spiral counts survived independent displacements of whole spacings, moving to a neighbouring pair and then refusing rather than misleading. Displacements with a direction are harder on it in only one case. A head pressed to an aspect ratio of 2.25, spread at the rim by eighty per cent or sheared with a slope of 1.6 is still counted as its own pair or its neighbour, and recovered inside its interval. A head twisted — each organ turned about the centre in proportion to its radius — is not: past a turn of the rim of about a radian and a half the counts leave their sequence and the recovered angle misses, by up to sixty-one degrees, because a twist changes the angle between one organ and the next. The round trip is not fooled. It is reporting the angle the twisted head has.

lattices · Recovery

The count sees the twist first

A twisted head recovers a changed divergence, and the check proposed for it was two annuli: the twist's extra angle falls with radius, so an inner and an outer annulus should disagree. Read on golden heads of 900, 2,400 and 9,000 organs, they never do in time. Their intervals separate at eight radians on 900 organs and never on the larger heads, always after the ordinary reading has been misled — from six radians on 900 organs and from two on 2,400 and 9,000. What catches the twist first, at every size and on every seed, is the count: at half a radian to three quarters some band stops returning two consecutive Fibonacci numbers — 34 and 89, 89 and 233 — which no untwisted golden head, clean or displaced, ever does.

lattices · Recovery

Where a handover sits

Inside every rung there is a rise at which the two contact steps change places, so that the shorter hop belongs to the other family below it. Six of the eight rungs on this ladder have one, each has exactly one, and every one of them sits in the coarse half.

cylinder · Rung interior

The family that lost a member

The offset rule restated in the arrangement predicts that the chain whose organ was taken is the chain that breaks. Scored on the nine offsets where the question can be asked, it is right none of the time and its opposite is right all nine.

emergence · Ablation

The organ that was nobody's neighbour

Twenty-one of the thirty wrecked offsets remove an organ that lies on neither contact chain through the tip. The reading that explains the other nine has nothing to say about them, and the honest thing is to say so rather than to widen the definition until it does.

emergence · Ablation

A stem too fine to settle

Below a rise of about four thousandths the counter stops returning contact families and starts returning pairs like 2/13 and 13/24. Lengthening the stem does not fix it. That is a ceiling on every sweep this collection runs up the ladder, and it has never been written down.

emergence · Noise colour

What a count cannot decide

A spiral count is the measurement this whole subject is built on, and it is deliberately blind to everything that varies inside a rung. Four results this collection now holds are results about that blindness rather than about the arrangements.

emergence · Counting

The organ that moved furthest

A reading that works on nine of thirty rows needs a reference organ, and the obvious repair is to measure one rather than to choose it. Measured, the disturbance turns out to have no far edge at all, so there is no organ that moved furthest in any sense the reading can use — and the generalisation that does work needs no reference organ.

emergence · Ablation

Named alongside it

The objects these essays reach for when they reach for this one.

Divergence angleMeasurementRiseParastichy pairLattice offsetCylinderLatticeHonest limitsNearest neighbourThe placement ruleClaim testingLadder

All concepts