Phyllotaxis
the spirals are counted, not admired
Concept

Rate — where it appears

How fast the rise falls as a stem grows, measured in organs per rung. It decides whether a pattern has time to change branch, and below a threshold a stem keeps the lattice it started on however long it is grown.

Named by 13 essays across 4 fields — each of them below, with the objects they name alongside it.

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Turing's final work was on phyllotaxis and it was unpublished when he died.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 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 48 cells with a carrier that pumps auxin up the gradient. Each short line is one cell's polarisation — the neighbour it pumps towards, which is always the richer one. 10 peaks come out, at a contrast of 93%, from a start that was uniform to within 6%.

A pump that works uphill

The mechanism that actually has molecular support behind it does not use a diffusing inhibitor at all. Cells move auxin towards whichever neighbour already has more of it, which is the opposite of what transport is supposed to do — and it produces a spacing from a field that started uniform to within six per cent.

mechanism · Auxin 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 What becomes of a seeded branch at 65 nodes per rung. Each bar is 10 runs at one amplitude, divided by what the blind counter found at the top of the stem. With no noise this rate ends on the Lucas branch. Placement noise displaces the node after the rule has chosen; field noise perturbs the energy the rule chooses over. Across 160 runs, 1 reached the Fibonacci branch with the lattice intact.

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 Only noise that arrives before the choice can change what is chosen. Intact runs only, from the whole amplitude sweep. Placement noise displaces the node after the rule has picked an azimuth: 44 runs, none of which changed branch at any amplitude that left a lattice. Field noise perturbs the energy profile the rule picks over, so it can move the minimum into a neighbouring gap: 1 of 66 did.

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 Both statistics, on the same stems, at a rise of 0.005. Five seeded stems at each disturbance, held at a fixed rise. Bars are how many returned the pair the position counter finds; open portions are refusals. The pair comes out from 0.1 to 0.25, and across that whole range the lag-one correlation of the same sequences is -0.33, -0.58, -0.59 — decisive, negative and flat. There is no trade between the two: one stem supplies both. Below the window the sequence has locked onto the sampling grid and is a cycle rather than a sample; above it there is no lattice left, at 117° of scatter.

What a refusal does not say

The readout can decline for four different reasons — too quiet, too disturbed, too fast, or a window in the wrong place — and a stem that returns nothing does not say which. That is the third time this thread has failed to close the mixture problem, and the first time the failure has a shape.

wrong · Survey spec Both ends of the window are silent, and a ruler tells them apart. The scatter recorded on stems at 400 nodes per rung, against the disturbance that produced it, with the stems that returned no reading at all marked as open. Silence at the quiet end comes with a scatter of 0.38 and 0.44°, which any botanist would call an orderly plant; silence at the disturbed end comes with 56°, which nobody would call a pattern. The two refusals look identical in the instrument's output and are three orders of magnitude apart in a quantity measured with a protractor.

A refusal with a reason

Three note left with the work running have recorded that a refusal has four causes and the sequence separates none of them. With a second window and a protractor, three are separated: silence at 0.38° of scatter is a quiet plant, silence at 56° is a disorderly one, and agreement certifies the rate. The fourth survives, and so does a worse discovery — agreement is not correctness.

wrong · Two windows Which rates keep a Lucas seed, on stems of one, two and three organs a whorl, placed by whorls a rung. Every rate each configuration was grown at, drawn at the number of whorls the stem places between one transition and the next: a dark cell keeps the Lucas ladder to the end of the rise, a light cell gives it up. 1 a whorl, 512: kept to 86.6 whorls a rung, lost from 87.6; 1 a whorl, 256: kept to 86.6 whorls a rung, lost from 87.6; 1 a whorl, 1024: kept to 86.6 whorls a rung, lost from 87.6; 2 a whorl, 1024: kept to 87.6 whorls a rung, lost from 88.1; 2 a whorl, 1024, imposed: kept to 87.6 whorls a rung, lost from 88.1; 2 a whorl, 512: kept to 87.6 whorls a rung, lost from 88.5; 2 a whorl, 2048: kept to 87.6 whorls a rung, lost from 88.5; 3 a whorl, 1536: kept to 87.9 whorls a rung, lost from 88.5; 3 a whorl, 1536, imposed: kept to 87.9 whorls a rung, lost from 88.5; 3 a whorl, 768: kept to 86.6 whorls a rung, lost from 88.2. On this axis every edge falls within about two whorls a rung of every other.

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 window that fits inside a rung. Stems that climb the ladder at four rates, read over a window at the fine end. The condition is a ratio: the window has to be shorter than a rung. 250 internodes at 130 per rung is 1.92 rungs and agrees on 0 of 3; 400 internodes at 130 per rung is 3.08 rungs and agrees on 0 of 3; 250 internodes at 260 per rung is 0.96 rungs and agrees on 3 of 3; 400 internodes at 260 per rung is 1.54 rungs and agrees on 1 of 3; 250 internodes at 520 per rung is 0.48 rungs and agrees on 2 of 3; 400 internodes at 520 per rung is 0.77 rungs and agrees on 3 of 3; 250 internodes at 1040 per rung is 0.24 rungs and agrees on 3 of 3; 400 internodes at 1040 per rung is 0.38 rungs and agrees on 3 of 3. Read over the whole stem instead, every rate returns nothing — 0 of 3, 0 of 3, 0 of 3, 0 of 3 — because the quantity the comb is periodic in changes as the pattern climbs.

A window inside a rung

A stem that climbs the ladder has no comb in it at any rate, because the quantity the comb is periodic in changes as it goes. Read a window instead and it comes back, on one condition: the window has to be shorter than a rung — which makes the shoot's rate the thing that decides whether a plant can be asked.

emergence · Rising Two windows on a shoot at 400 nodes per rung. A stem grown at 400 nodes to the rung with a disturbance of 0.25, its rise falling from 0.4 to 0.004 over 1914 nodes. The upper window is the last 250 internodes — where a count would be made on a real plant — and the lower is the same length shifted down 125. The upper reads 8/13; the lower reads 8/13. The verdict is agree, and the window holds 0.63 of a rung.

Two windows on one stem

A pair read off a climbing shoot can only be read through a window, and a window can straddle a transition. Read a second window half a length lower and the outcomes fall into four kinds — and agreement between them never happens on a shoot whose rung is shorter than the window, which turns the most awkward of the four refusal causes into something a reading can certify.

emergence · Two windows

Named alongside it

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

Divergence angleRiseBranchHonest limitsRungAutocorrelationIdentifiabilityMeasurementNoiseThe placement ruleSpecimenAuxin

All concepts