Concept

Branch — where it appears

One of the families of divergence angles a placement rule can settle on, named by the sequence its counts follow. The Fibonacci branch is the common one and the Lucas branch is the usual exception, and which a stem takes is decided by its seed and its rate.

Named by 37 essays across 6 fields — each of them below, with the objects they name alongside it.

The rule, 26 steps in, at a growth of 0.40. The next primordium goes where the repulsion is least — the marked minimum at 216°. Nothing in the rule refers to any particular angle.

The angle is an output

137.5° is not a constant of nature. It is where a rule settles — a rule that places each new element as far as it can from the ones already there, contains no reference to the golden ratio, and reaches the same answer from starting angles a hundred and sixty degrees apart.

emergence · Attractor
A branching tree in which every junction obeys the cube law. r₀³ = r₁³ + r₂³ at all 63 junctions, to 2e-16. The widths in the drawing are the radii the law gives, not widths chosen to look right.

The cube law

A branching network built to move fluid for the least work obeys one relation at every junction — the cube of the parent radius equals the sum of the cubes of the daughters. It is a minimisation result, it is checkable on a real tree, and it is the rare biological rule with a derivation.

branching · Murray
The spiral counts four different divergence angles produce. Fibonacci counts come from one angle. The Lucas angle — which the same dynamical model reaches on a different branch — gives 47 and 76, and neither number is a Fibonacci number.

Fibonacci is a branch, not a law

Fibonacci counts come from one branch of the model. The Lucas branch — which the same rule reaches at a different growth rate — gives 47 and 76, and neither is a Fibonacci number. The sequence is a consequence of an angle rather than a property of plants.

wrong · Fibbranch
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 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
What the model settles on, against how fast the meristem grows. A broad golden branch, a transition, and then the two-whorl regime at exactly half a turn. 9 of 27 converged settings land within 4° of the golden angle; 15 land more than 20° away.

The bifurcation diagram

Sweep the one parameter of the rule and the settled angle traces a diagram — a broad golden branch, a transition, and a two-whorl regime at exactly half a turn. The famous constant is one branch of it, which is a more useful thing to know than the constant.

emergence · Bifurcation
The version of the claim that does survive measurement. The golden angle scores 0.4377, against 0.3462 for the best of 938 other angles sampled. The dashed line is Hurwitz's 1/√5, which no number can exceed.

The claim that survives

Of the three famous assertions about this subject, one is out by a factor of two, one is true of a branch rather than of plants, and one is right — in a sharper form than the version usually told, and about arithmetic rather than about packing.

wrong · Hurwitz
An L-system after 4 rewrites of two rules. X → F[+X]F[-X]+X and F → FF, walked by a turtle turning 22.5°. 130 segments, and not one of them knows anything about light, water or auxin.

L-systems describe, they do not explain

Two rewriting rules and a turtle produce something indistinguishable from a plant, and there is no plant in it — no light, no water, no auxin, no mechanics. That is worth demonstrating precisely because the output is so convincing.

branching · Lsystem
The spiral counts, band by band, in one head. The same flower gives 13/21, 21/34, 34/55, 55/89 at different radii. A caption saying "34 and 55 spirals" is a statement about one annulus.

The counts change with radius

The same head gives 13 and 21 near the centre, 21 and 34 further out, 34 and 55 beyond that, and 55 and 89 at the rim. The transitions are at computable radii, and a photograph captioned with one pair is a statement about one annulus rather than about a flower.

lattices · Transitions
Two runs of the same rule from unrelated starting angles. Both settle at 137.0°, within 0.5° of the golden angle, from seeds 166° apart.

Droplets with no biology in them

Douady and Couder dripped magnetised ferrofluid into a dish of silicone oil, and got spiral phyllotaxis with Fibonacci parastichy numbers out of a system containing no cells, no genes and no plant. That is the strongest evidence the pattern is physics — and the clearest warning about what a model can claim.

emergence · Analogue
A golden spiral and a nautilus spiral over 2.5 turns, from the same start. After 2.5 turns the golden curve is 7× larger. The growth factors are 6.85 and 3.2, a factor of 2.14 apart.

The nautilus question

A golden spiral grows by 6.854 per turn. Measured nautilus sections give about 3.2. That is a factor of 2.14 — not a rounding error, not an artefact of where the centre is assumed to be, and not close.

shells · Nautilus
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 this implementation stops converging. Below about G = 0.18 the settled angle wanders over 113° however long the run. That is the model's limit, not a fact about plants.

Where the model stops

Below a growth parameter of about 0.18 this implementation does not converge — the settled angle wanders over a hundred degrees however long the run. That is the range where the literature says the interesting behaviour lives, and it is worth a figure rather than a quietly chosen axis.

emergence · Modellimit
What a divergence picked at random gives, at a rise of 0.100. Fibonacci pairs take 59.6% of the circle at this rise, and the share falls as the rise does. The claim that Fibonacci counts are what nature "prefers" needs the preference to come from somewhere, and it is not from the geometry being generous.

How often is it Fibonacci

The claim that plant spirals come in consecutive Fibonacci numbers is stated as a near-universal. Asked of the geometry, the answer collapses with scale — at a coarse rise 67% of divergences give Fibonacci pairs, and at a fine one 15%, with whorled and unnamed pairs taking the rest.

wrong · Frequency
A tree built at an exponent of 3, measured band by band. Six bands of daughter ratio, 300 junctions in each, all from trees built at an exponent of exactly 3 with the same 2% measurement error. The median implied exponent falls from 3.00 at an even fork to 1.58 at a twig, and the share of junctions that have any exponent at all falls from 100% to 53% over the same range. The rule across the top is a single fit over 200 junctions spanning the whole range: 2.85. A mixed sample is safe because least squares already weights by leverage — 42% of it sits in the most symmetric band and 0.06% in the twigs. The sample that is not safe is the one a person can reach.

The band decides the answer

A fit over a whole tree's junctions returns the exponent the tree was built at, even though most of its junctions are from bands that on their own return 1.6. Least squares is already weighting by leverage. The dangerous sample is not the mixed one — it is the one a person can reach.

branching · Exponent
Two paths down the same tree. Both start at the same first fork. Keeping the larger family every time reaches 137.473°; one different choice reaches 99.549°. Neither angle is in the arithmetic — both are limits of a path.

The tree and the attractor

The dynamical model settles on the golden angle over a range of one parameter and on the Lucas angle outside it, and it cannot reach the low-growth end at all. The lattice tree reaches everywhere, has no dynamics in it, and produces the same two angles as limits of two paths. Two routes, one pair of numbers.

emergence · Branch tree
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 stem grown at 42 nodes per rung. 167 nodes, each placed where the repulsion from the ones below it was least, with the rise falling from 0.2 to 0.0045. Counted blind in a sliding window the pattern walks 1/2 → 2/3 → 3/5 → 5/8, and the marks are where its answer changed.

A pattern with a rate

Every lattice in the essays before this one is a static object indexed by a parameter, and a plant is not. Put the rise on a clock, place each node where the repulsion from the ones below it is least, and the object that comes out has a history — which is the first thing here that could disagree with the ladder.

emergence · Rising
The 2-jugate forks converge on 68.7539°. Every fork sits at a rational divergence, with denominator 4(m² + mn + n²) — 10/28, 30/76, 74/196 and so on. The limit is 68.7539°, which is 137.5078 divided by 2, and it is at none of them.

Half the golden angle

The forks of the van Iterson tree converge on 137.5078° and sit at none of them. Divide the whole tree by two and the same thing happens at 68.7539° — which is where teasel is, and where a bijugate sunflower counted 42 and 68 has to be.

lattices · Jugate limit
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 lag between two rates, measured against the rate. Each dot is one rate: the mean gap between where the grown pattern changed its count and where the static ladder puts that transition, in rungs. Over rates from 9 to 135 nodes per rung the worst is 0.087 of a rung. A lag of one rung would put a dot on the top line.

The lag that is not there

A pattern built out of its own history should hold its old parastichy pair past the point where a fresh lattice would have changed, and the gap should grow as the shoot is hurried. Over a fifteenfold range of rate it does not — every transition lands within a tenth of a rung of where the static ladder puts it.

emergence · Tracking
One seed, two rates, two ladders. Both stems begin as forty nodes of Lucas lattice at a rise of 0.12. At 65 nodes per rung the divergence stays at 99.5° and the counts walk 1/3 → 3/4 → 4/7 → 7/11. At 131 it leaves for 137.7° and walks 1/3 → 2/3 → 3/5 → 5/8 → 8/13 instead.

The rate decides the branch

Forty nodes of Lucas lattice, carried down to the same fine rise twice. Hurried, the pattern holds the Lucas ladder through three more forks at 99.5°. Given room, it abandons it at the first fork it reaches and walks to 8/13 at 137.5°. Same seed, same rise, two ladders — with a threshold between them at about ninety nodes per rung.

emergence · Branch choice
Fibonacci at a rise of 4.8e-3, asked two ways. Choose a divergence at random and a static lattice at this rise gives a consecutive Fibonacci pair 10.8 per cent of the time. Start coarse at a divergence nobody chose, grow the stem down to the same rise, and it is 100 per cent of 16 runs. The geometry is not generous; continuity is.

Continuity from a coarse start

At a fine rise, one divergence in seven gives a Fibonacci pair. Grow a stem from a coarse start at a divergence nobody chose, down to that same rise, and sixteen runs out of sixteen end on 8/13. The earlier work's interpretation was that continuity does the work; this is the measurement it never had.

emergence · Continuity
The angles against the positions, rise by rise. five 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 0.01 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. At 0.008 the counter says 5/8 and the angles agree on 2 of 5, refusing 3. The two instruments share no code path: one is given a list of angles, the other a list of coordinates.

The angles name the branch

Seed the same rule at the Lucas angle and the readout returns 4 and 7, then 7 and 11 — the pairs the position counter finds, and not Fibonacci numbers. So a list of divergence angles carries not only how many spirals there are but which family of ladders the plant is on.

lattices · Counting

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

Noise is not a slow rate

A stem seeded on the Lucas branch keeps it below ninety nodes per rung and abandons it above — which invites the objection that a real apex's fluctuations would knock it off regardless. Measured across a hundred and sixty runs of two independent kinds of noise, one escapes, at the amplitude where the pattern is already coming apart.

emergence · Noise amplitude

Where the noise gets in

Ninety runs of noise applied after the rule has chosen, and not one changes branch. Fifty-six of noise applied to the choice itself, and one does. Only a disturbance upstream of the decision can restructure which nodes are neighbours of which — which is what a branch is.

emergence · Noise entry

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

A cube law with a lever arm

Murray's exponent of three comes from moving fluid for the least work, and Da Vinci's two has had no derivation here, only the name of the mechanical answer. Size every branch so that the same wind on every tip bends it to the same stress, and a junction conserves r to the power 3/(1 + log₂(1/λ)), where λ is how much shorter each branch is than its parent. A crown that fills a plane gives exactly two; halving lengths gives one and a half; no shortening gives three. Murray's flow rule gives three at every λ, so the lengths of a tree's branches say which mechanism sized it.

branching · Murray

A Lucas seed counts whorls

An ordinary stem seeded on the Lucas lattice keeps that ladder when its rise falls fast and gives it up when it falls slowly, with the edge near ninety nodes a rung. A stem that grows two organs a whorl is an ordinary stem folded twice round, so the identity predicts its edge — once it says whether the edge counts placements or whorls. Grown across the edge, stems of one, two and three organs a whorl keep a Lucas seed to 86.6, 87.6 and 87.9 whorls a rung: one number to within a per cent in whorls, and one, two and three times as far out in organs. No grid moves it and imposing exact whorls moves it not at all, even where free trijugate whorls come apart completely as the seed is lost.

lattices · Jugate limit

A count set by a delay

An L-system describes a plant and forbids nothing, because none of its parameters is anything a plant has. One branching grammar is the exception: a mature apex makes a new bud every season, and a bud waits d seasons before it branches. Its counts grow at the root of x^(d+1) = x^d + 1, a delay of one season gives Fibonacci's numbers and nothing else does, and the fourth count already separates a one-season wait from every longer one. So a Fibonacci count in a branching plant is a measurement of how long its buds wait. It is also a fragile one: if one bud in ten waits two seasons instead, eleven counts in a row come out Fibonacci's three times in a thousand.

branching · Lsystem

A count that loses its growing points

The branching grammar behind the Fibonacci claim has no deaths in it, and a stem that loses shoots is the common case. Giving every growing point a chance q of dying each season leaves the counts a linear recurrence and does exactly one thing to it: the growth rate becomes the deathless root multiplied by 1 − q, at every delay and every death chance, to the last bit a double holds. So each waiting time has a death chance above which its lineage shrinks — a half with no wait, 0.3820 at one season, 0.2451 at four — and a longer wait tolerates less. What does not survive is the count itself: a plant losing one growing point in ten a season shows eight Fibonacci counts in a row one time in ten thousand, against one time in eight for a bud that occasionally waits an extra season.

branching · Lsystem

What a scar is worth

Counting the scars a dead shoot leaves does not put a branching count back on the sequence it would have had. A scar records a growing point and a dead growing point takes every branch it would have made, so living points plus scars reach 39.2 per cent of the deathless count after twenty seasons at one death in twenty, and 1.9 per cent at one in five — falling without limit rather than closing. What the scars restore is the other number. Scars per living point settle at q/(x − 1) exactly, so a rate with a scar share beside it recovers the death chance and then the waiting time, where a rate alone is reached by a one-season wait losing a tenth, a two-season wait losing 0.64 per cent and no wait at all losing 27.2 per cent.

branching · Lsystem

Seven rises and two seeds

One organ removed from a stem is felt out to the larger of its two spiral counts. Every test of that has confounded the count with the rise, because on one branch the two move together. Grow a second branch beside the first at the same rise and they come apart — and doing it at seven rises turns a matched pair into a design whose last column changes hands four times.

cylinder · Branch choice

The front that reads one short

Eleven cells of a fourteen-cell design put the boundary exactly at the larger spiral count. Three put it one offset earlier, and the tempting move is to lower the threshold until all fourteen agree. Measured instead of tuned, the three turn out to be the three cells nearest below their own rung's boundary — and the last offset of a front is weak because it has only just arrived.

cylinder · Branch choice

The hole on the other branch

Near a transition, the run of offsets a stem notices stops being a run: there is quiet past the front and then one isolated offset, felt as hard as anything inside it. Where that offset sits was pinned down on Fibonacci lattices, where the numbers to check it against are 5, 8 and 13. On the Lucas branch they are 4, 7 and 11 — and the rule holds there too.

cylinder · Branch choice

The branch is what is left

Four accounts of why one band's cuts change what they keep and another's do not were written down before a third band was cut. Three of them are now wrong on a band each, and the survivor is the one with no mechanism behind it.

cylinder · Rung interior

Named alongside it

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

FibonacciRiseDivergence angleLadderHonest limitsRepulsionClaim testingMeasurementTransitionsAttractorMeristemParastichy pair

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