Concept

Cylinder — where it appears

The surface almost all real phyllotaxis happens on, with an azimuth and a height rather than a radius. A disc is a cylinder whose rise falls as the square of the reciprocal radius, which is what lets one model predict where a seed head's counts change.

Named by 31 essays across 7 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
Which patterns grow on a ring of circumference 1.40. Modes 4 to 15 have positive growth rates and mode 8 is fastest. Integrating the full equations from a disordered start gives 8 peaks.

Turing's last problem

Turing's final work was on phyllotaxis and it was unpublished when he died. Its core is a ring of cells and two diffusing substances, and the thing it does is select a number of peaks — which can be predicted from the equations before anything is integrated, and then counted from what the integration produces.

mechanism · Reaction–diffusion
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 angle between the first two peaks, for six starting disorders. The same equations, the same ring, six different starting perturbations: 177°, 47°, 109°, 151°, 47°, 133°. A spiral needs the angle between successive elements to be the same one each time, and a stationary ring does not supply that.

A ring cannot make a spiral

The peaks on a Turing ring do not all appear at once — there is a first and a second. But which two lead is decided by the starting disorder, so the angle between them comes out at 177°, then 47°, then 109°, then 151°. A divergence angle is a relationship that repeats, and this one does not.

mechanism · Ring limit
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
The exponent fitted from the junctions, rather than assumed. Sweeping k and asking where r₀ᵏ = Σ rᵢᵏ holds best gives 3.000 — Murray's 3, recovered rather than imposed.

Fitting the exponent

Assuming the exponent is three and reporting the error says how far the data is from that assumption. Fitting the exponent and reporting what it comes out as says what the network is doing — and an estimator has to be shown returning something other than three, or it is not a fit.

branching · Exponent
Closest pair across 120–155° at 400 organs, read both ways. On the interior's scale the golden angle reads 0.9027 and ranks 1st of 72, against 0.9026 for the best grid angle at 137.5°, with the window running from 0.0668 to 0.9026; counting the rim's cells the golden angle reads 0.7076 and ranks 2nd of 72, against 0.7129 for the best grid angle at 137.5°, with the window running from 0.0331 to 0.7129. The dashed upright is the golden angle, which a grid of decimal degrees never lands on and which is therefore read separately.

Packing, measured four ways

The claim is that the golden angle packs best, and it is measurable. Read on the interior of a head, the two criteria about distance put the golden angle first among the angles near it and the two about cells are won by rational angles — which makes the claim half right, and makes the right half a statement about a class of angles. An earlier reading of the same four criteria, divided by the cells at the head's edge, said the opposite.

tissue · Packing
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
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
The plane of stems: divergence across, rise up. Each shade is one parastichy pair. The marked points are the lattices where three families are equally short — the forks — and the Fibonacci ones run up the middle towards 137.51°.

The forks are exact

Where a stem's pattern has to choose between two futures, three spiral families are equally short and the lattice is exactly equilateral. A numerical solver found those points; the numbers it returned turned out to be rational, and chasing that gave a closed form — including the fact that every fork sits at a rational divergence, and the golden angle at none of them.

cylinder · Forks
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
The plane of stems: divergence across, rise up. Each shade is one parastichy pair. The marked points are the lattices where three families are equally short — the forks — and the Fibonacci ones run up the middle towards 137.51°.

The angle is not the object

Every popular account of phyllotaxis is organised around a number. After a round of work spent on stems, forks and frequencies, the number looks like the wrong thing to organise an account around — it is the limit of one path through a branching structure, it is at no fork, and a plant that has it got there by not jumping.

wrong · The tree not the angle
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
The rise falls as one over the distance from the apex. A straight line of slope −1 on log axes. The dashed horizontals are the transition rises of the cylinder's ladder, computed with no cone anywhere in them; where they cross, the count changes. Consecutive crossings are 2.62, 2.62, 2.62, 2.62 apart — φ² is 2.618.

Transitions a factor of φ² apart

The ladder's rungs are a factor of 1/φ² apart in rise. A disc's rise falls as one over radius squared and a cone's as one over distance, so the same rungs land a factor of φ apart on a seed head and a factor of φ² apart on a cone — measured, on both, by a counter that has never heard of either.

cylinder · Cone transitions
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
Every open question here needs under 34 specimens. The sample size at which each comparison reaches 90 per cent power at a 5 per cent false-positive rate, from the exact binomial rather than a normal approximation. The census question — do plants show consecutive Fibonacci pairs far more often than the geometry does — needs 4: 14.7% is the share of divergence angles giving a consecutive Fibonacci pair at a fine rise; 90% is what a grown history gives.

The survey this site cannot do

Four rounds of asking for a dataset, and it is still not here. What the work here can do instead is specify it — the fields, the sampling, the sizes, and which of this collection's claims each one would settle. Two of the four fields asked for turn out to be worth less than the asking implied, and one was never asked for at all.

wrong · Survey spec
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
Which neighbours decide where an element goes. Each line is one exponent: how much each shell of neighbours makes the energy profile vary around the circumference, divided by what the nearest shell contributes. At p = 0.5 the nearest shell leads the next by a factor of 1.1 and a node is placed against the whole neighbourhood at once. At p = 3 it leads by 9.7e+3, and a node is placed against its immediate neighbours — which is what a lattice is.

How far a primordium reaches

The placement rule's repulsion falls as an inverse cube because that is what two magnetised droplets do, and nothing about a plant supplies the exponent. Asking what it controls produced one tidy wrong answer and one measured right one — and the difference between them is the difference between a total and a variation.

emergence · The range of the interaction
The memory of a divergence sequence, at 0.75° of scatter. With no noise at all the lag-one correlation is 0.54: the rule corrects itself, so a lattice arrives with a memory in it. Matched at the same recorded scatter, placement noise leaves -0.10, jostle noise leaves 0.66, field noise leaves 0.47. The band is ±0.13, which is what an uncorrelated sequence of this length gives.

The sequence has a memory

Every measurement this collection has made of a stem's divergence angles throws the order away. A spread is invariant to shuffling. Put the angles back in order and there is a large correlation between one and the next — 0.54 with no noise at all — which is the rule correcting itself, and which nothing had looked at.

cylinder · Sequence
A rule too long-ranged makes no pattern; every shorter one makes the same pattern. Each dot is 4 runs from a coarse start at one exponent, separated by 0.2° of placement noise. Below p ≈ 1.1 the divergence scatters by tens of degrees, which is what an arbitrary sequence gives. From p = 1.25 to p = 8 — a sixfold range — every run ends on 8/13 with the scatter between 0.75° and 1.06°.

The exponent that barely matters

An unchecked claim, repeated since the first essays, held that the repulsion's falloff exponent hardly changes the answer. It could not be tested, because the function that would have taken it never passed one down. Tested at last, it is true on a disc — by three and a half degrees across a sixteenfold range — and on a stem it decides whether there is a pattern at all.

emergence · Falloff exponent
What the sequence sees that the scatter cannot. Each point is an ensemble at one amplitude, placed at the scatter it produces. A stem at three quarters of a degree of scatter has a lag-one correlation near zero if its noise arrived after the primordium was placed, and near 0.7 if it arrived before — and no measurement of the scatter can tell those apart. The separation closes above about a degree, because what the other two kinds preserve is the correlation of a lattice.

What one angle says about the next

A tenth of a degree of placement noise moves a stem's divergence scatter from 0.50° to 0.62°, which nobody would report. It takes the correlation between consecutive angles from 0.54 to below zero. The other two kinds of noise, at scatters where no measurement can separate them, leave it at 0.6.

cylinder · Sequence
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

The memory was the rise

The earlier work measured a lag-one correlation of 0.54 in a noiseless divergence sequence and called it the sequence's own memory. Hold the rise fixed and there is no sequence at all — every angle identical — and under a disturbance the correlation is negative. The 0.54 belongs to the pattern chasing an equilibrium that is moving under it.

cylinder · Sequence

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

How many organs a pair needs

A count taken over too few organs does not fail. It returns the rung below, which is a perfectly good pair, and nothing anywhere says so. The window that avoids it is not a constant but the counter's own arithmetic, and 384 settled runs sit exactly where that arithmetic puts them.

lattices · Read window

The neighbourhood was already settled

The earlier work explained a small difference between two kinds of noise by saying a jostle is diluted among some thirty neighbours. Sweep the neighbourhood sixfold and the difference does not move — because past four spacings the rule builds the identical lattice, internode for internode. There was nothing to dilute.

mechanism · Noise entry

Two thirds of a cell

The founding claim of this field is that the six sides Euler forces are the spiral families. Measured against the tessellation it names two thirds of a cell's walls exactly, in every band of a head and at every rise of a stem, and the missing third is the same third everywhere.

tissue · Neighbour definition

A shoot too fast to remember

Sweep the rate at which a stem climbs the ladder and the correlation between one divergence and the next changes sign — negative below about fifty-five nodes per rung, positive above it, with the flip inside one step of the grid. The instrument the earlier work proposed is unavailable on a fast shoot, and nothing said so.

emergence · Rising

The third family

On a seed head no threshold makes the counted contacts and the shared cell walls the same relation. On a stem they are the same relation exactly, at every one of a hundred and eleven rises and to three decimal places of nothing, provided the contact cut keeps three families where a count keeps two.

tissue · Neighbour definition

Named alongside it

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

RiseDivergence angleParastichyMeasurementLadderThe placement ruleEnsembleFibonacciNoiseAutocorrelationEquilibriumParastichy pair

All concepts