Concept

Transitions — where it appears

The rises at which a lattice changes from one parastichy pair to the next. They are computed from the geometry as the rises where two lattice vectors have equal length, so a prediction about where a seed head's counts change is exact rather than fitted.

Named by 32 essays across 5 fields — each of them below, with the objects they name alongside it.

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 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
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
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
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 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 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
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
What is visible in the outer part of a 4000-element organ. Both surfaces have the same ladder in element number — the rise is 1/(2πi·flare) on a cone and 1/(4πi) on a disc, and c and the internode step both cancel. What differs is where the elements are. Counting outside 50 per cent of the extent, a cone shows 1 change and a disc 2, because half a cone's length holds half its elements and half a disc's radius holds three quarters of them.

Why a cone can be counted once

A pineapple is described as 8 and 13 and the description holds. A sunflower is described as 34 and 55 and the description is a statement about one annulus. Both organs have the same ladder in element number — what differs is where an organ puts its elements.

cylinder · Organ size
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 exponent fitted to an organ that has 4 of them. Each dot is one step between consecutive rings, reporting 2 ln φ / ln(s′/s) — the exponent that step would have if the organ had one. They run from 1.980 to 1.697. The line is what a single fit returns, 1.891, which is their harmonic mean of 1.880 and sits below their plain average of 1.887.

What one exponent reports

Fit a single shape exponent to an organ that has four of them and it returns a real quantity — the harmonic mean of what its individual steps report. Harmonic means sit below arithmetic ones, so the fit understates, systematically, in a known direction, and invisibly.

cylinder · Fitted exponent
At 1.0 per cent on each ring, 5 rings show the drift. The gaps between consecutive rings are 1.626, 1.631, 1.657, 1.763. Two rings give one gap and no way to disagree with itself; three give two gaps and a fit with nothing left over. The question is how many gaps it takes for their spread to exceed what the measuring error can explain, and the answer depends on the error as much as on the organ.

How much of a cone to measure

Two rings cannot show a varying exponent — not with difficulty, but in principle, because one gap determines one exponent with nothing left to disagree. Four or five can, if each is found to within a per cent. At three per cent this specimen cannot be told from a power law however many of its rings are recorded.

cylinder · Ring identifiability
The defect rings are not where the counts change — they are √φ further out. A logarithmic radius axis for a golden, 137.508° head. The lower marks are the radii at which the counted parastichy pair changes, where the two shortest lattice vectors change places; the upper marks are the radii at which a cell's neighbours change, where the third-shortest does. They interleave, and the ratio of each ring to the transition inside it is 1.2715, 1.2723, 1.2723, 1.2719, 1.2719, 1.2723 against √φ = 1.27202. Consecutive rungs are a factor of φ apart in radius and √φ is their geometric midpoint, so a defect ring sits exactly halfway between two parastichy transitions. Anyone looking for the defect line at the radius where the counts change will not find it there.

The rings are not the transitions

A seed head has two ladders on it — the radii where the counted parastichy pair changes, and the radii where the exceptional cells sit — and the obvious guess is that they are the same ladder. They are not: the second sits a factor of the square root of phi outside the first at every rung of two different divergence ladders, which is exactly halfway between two consecutive transitions.

tissue · Topological charge
The pairs grown stems of one, two, three and four organs a whorl count as their rise falls, against the folded rise. Four stems grown by the same rule over the same range of folded rise, 0.4 to 0.0012 — the rise between whorls times the jugacy, which puts every jugacy on one lattice axis — each whorl folded to one node and counted in a sliding window. With 1 organ a whorl the counted pairs run 1/2, 2/3, 3/5, 5/8, 8/13, 13/21; with 2 organs a whorl the counted pairs run 2/4, 4/6, 6/10, 10/16, 16/26, 26/42; with 3 organs a whorl the counted pairs run 3/6, 6/9, 9/15, 15/24, 24/39, 39/63; with 4 organs a whorl the counted pairs run 4/8, 8/12, 12/20, 20/32, 32/52, 52/84. Every jugacy passes the same transitions at the same folded rises, with its pairs multiplied by its jugacy.

A grown stem halves its ladder

A bijugate stem at 68.754° was derived to pass the ordinary transitions at half their rises, because a lattice wrapped twice round is the ordinary lattice at twice the rise. No bijugate stem had been grown through them. Grown by the same rule that grows an ordinary shoot, stems of two, three and four organs a whorl walk the ordinary ladder with every pair multiplied by the jugacy, change pair at the ordinary rise divided by the jugacy between whorls — and by its square per organ — lag behind the static ladder by the same hundredth of a rung, and settle within three hundredths of a degree of 137.5078 over the jugacy.

lattices · Jugate limit
The ratio is a U across every rung, and its floor is the number that was reported. The ratio of the second comb to the main comb, on five stems at each of 9 rises spanning two rungs, against the ladder's own coordinate for where each rise sits inside its rung. Both rungs give the same shape: a floor of 0.71 and 0.79 about two thirds of the way up, climbing towards the transition at either end. The dashed line is a transported disturbance with no rule in it at 1.28, which does not vary with the rise at all — a kinematic lattice's angle sequence has no rise in it. Where the rule's curve crosses that line the two accounts are indistinguishable.

The survey loses its second outcome

The survey specification written earlier here names three results the survey could return, and the second — a ratio near or above 1.30, read as evidence against the placement rule — is the one that would have been worth publishing. It does not survive the measurements here. The ratio moves with where the plant sits between two transitions, and it moves again with the colour of the plant's own disturbance.

wrong · Survey spec
One cell's six walls, and the two the pair 34 and 55 does not name. The cell of primordium 225, at 50 per cent of the head's radius, with each of its six walls labelled by the index difference across it. Its own counting instrument returns 34 and 55, which names four of them; the two drawn warm are 21 and 21, a family the instrument ranked and discarded. Over the whole head that is 33.80 per cent of the union in dispute, and it is the same fraction in every band.

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
On a rung the response is a run of offsets; near a transition it has a hole. One row per rise, from 0.02 at the top to 0.005 at the bottom, and one column per offset: the organ one place back at the left, 14 places back at the right. A cell is filled where removing that organ moves the next organ by more than 2.5°, and empty where it does not. On a rung the filled cells are a run from one to the larger parastichy number — 8, 12, 13 at the pairs shown on the left. Between a rise of 0.02 and 0.008 the run ends at 5 and one more cell is filled at 7, with the offsets between them quiet to under a degree. That isolated column is one place inside the larger number of the pair the stem is climbing towards.

The ablation a plant would survive

The intervention proposed earlier returns a spiral count from a yes-or-no answer, needs no protractor, and was specified at one rise. Measured across the ladder it acquires three conditions a real experiment would have to meet — and one of them is that the plant must not be too coarsely patterned, or nothing will go wrong at all.

wrong · Survey spec
The ratio is a U across every rung, and its floor is the number that was reported. The ratio of the second comb to the main comb, on five stems at each of 15 rises spanning two rungs, against the ladder's own coordinate for where each rise sits inside its rung. Both rungs give the same shape: a floor of 0.71 and 0.79 about two thirds of the way up, climbing towards the transition at either end. The dashed line is a transported disturbance with no rule in it at 1.29, which does not vary with the rise at all — a kinematic lattice's angle sequence has no rise in it. Where the rule's curve crosses that line the two accounts are indistinguishable.

The ratio was the floor of a curve

One number was left standing between a placement rule and a transported disturbance, measured at one rise, with the explanation that the geometry there happens to favour the larger parastichy number. Swept across two rungs the number is a U — a floor of about 0.79 two thirds of the way up a rung, climbing past 2.8 as a transition approaches — and the geometry is flat exactly where the curve is steepest.

cylinder · Noise transport
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
A stem is a thread at six rises of the sweep, a ribbon at 23 and a surface at 111. The 140 rises of the sweep, split by how many index families the tessellation carries. On 6 rises above 0.5010 a node has 2 walls and the strip is a thread; on 23 between 0.2187 and 0.4833 it has 4 and is a ribbon; on 111 below 0.2109 it has 6 and is a surface. The comparison means something different in each: two families leave 33.333 per cent in dispute on the surface and 0.00 on the ribbon, and three families leave 0.000 and 33.333.

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
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

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 response with a hole in it

Removing an organ is felt out to the larger parastichy number and no further — that is the intervention's headline, and it holds in the middle of a rung. Swept towards a transition the run of felt offsets stops early and one lone offset past it comes alive, with three quiet organs in between. The lone offset is one place inside the count the stem is about to have.

emergence · Ablation

A stem coarse enough to cut

Below the 3/5 rung is a 2/3 rung, and it runs from a rise of 0.050 to 0.120. It is not a lattice across all of it: from 0.090 to 0.115 the divergence stops settling and sticks on exactly three eighths of a turn, wobbling by a degree and a half — while a counter goes on reporting 2/3 as though nothing had happened.

cylinder · Coarse rung

The organ that guards the second slot

An organ twelve places back is the furthest of any from where the next one goes, and removing it moves the next one by a whole divergence. The reason is that the rule's profile has two low points rather than one, the second is the slot after next, and that organ is holding it up. The comparison between what it holds up and how far behind it is decides the whole thing.

emergence · Ablation

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

Two lines that cross once

The divergence at which a lattice's two contact steps are exactly equal is a curve across each rung, computable from the geometry with nothing grown. The rule's own settled divergence is a second, shallower curve, and where they cross is where the step ordering changes hands.

cylinder · Rung interior

A wall and not a budget

Below a rise of about 0.005 this collection's stems stop settling onto a lattice, and the limit has been written up four times without anybody asking which kind of limit it is. Grown three times as long, the table is identical row for row: not one stem that failed to settle succeeds. The floor is a wall.

cylinder · Settling

Named alongside it

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

RiseLadderParastichy pairHonest limitsRungMeasurementDivergence angleBranchSpecimenArtefactCounting blindParastichy

All concepts