Phyllotaxis
the spirals are counted, not admired
Theme

The thread: The round trip

Build the pattern from a stated number, forget the number, recover it from the pattern alone, and compare. It works for the divergence angle and for the growth factor, and the agreement is the measurement.
A logarithmic spiral growing by 3.20× per turn. Fitting log r against angle on the drawn points returns 3.2000, and the model's worst residual is 9e-16 in log r. Shells and growth

Growth as a rule

A logarithmic spiral is not a shape somebody admired. It is what a thing grows into when it adds material at its opening without changing shape, its one parameter is how much it grows per turn, and that parameter can be recovered from any drawn curve to the last digit.

 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. The pattern itself

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.

 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. Branching and transport

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.

 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°. Stems and cones

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.

 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°. The pattern itself

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.

 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. The pattern itself

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.

 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°. Stems and cones

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.

 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. The pattern itself

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.

 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. The pattern itself

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.

 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 pattern itself

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.

 The two halves of a 3.2 spiral fitted about a centre 0.25 innermost radii off, towards 52°. A logarithmic spiral growing by 3.2 a turn over 4 turns, which does not change, split into an inner half and an outer half, each fitted about a centre displaced by 0.25 of the innermost radius towards 52°. The inner half returns 3.4193 and the outer half 3.2200, a split of −5.83 per cent. The panel on the right enlarges the first whorl, where the true centre and the assumed one can be told apart; across the whole spiral the displacement is 0.238 per cent of the outer radius. Shells and growth

A centre that invents a life history

The collection's advice for a shell that might have changed how it grew was to fit it twice, over different arcs, and compare. On a spiral that does not change at all, a centre displaced by a quarter of the innermost radius splits the two halves by 4.09 per cent — the split a genuine 8.35 per cent change from apex to aperture produces — in either sign, depending only on which way the centre is wrong. Point noise of the same size splits them by less than half as much, and averages away where the centre does not. The floor under the test is the centre, not the noise.

 A shell whose deposition law changes part way through, beside one that never does. Both panels are the same logarithmic spiral at 3.20 per turn, marked at equal intervals of time. On the left the animal holds a constant angular rate for the first 3.50 whorls and a constant length added after that; on the right it holds a constant length added throughout. The curves are identical to the last bit a double holds, because a curve records no clock at all. The counts per whorl are not: 56, 57, 56, 67, 191, 613, 1960 against the unchanged shell's, and the change is in where the marks crowd rather than in where the shell goes. Shells and growth

A shell that changed its law

An animal that grew as a juvenile under one deposition law and as an adult under another leaves a sequence of whorl ratios rather than one, and the sequence says where the change happened. The ratio across the change is a closed form that is neither law's — 6.72 between a length clock and an area clock at 3.2 per turn, exactly the average of 3.2 and 10.24 — and it is monotone in where inside its whorl the change sits, so it inverts. On a seven-whorl shell of 18,466 lines a change at 3.5 whorls comes back at 3.5001, in a band 0.027 whorls wide that holds the true position. The reading refuses a change in the outer three whorls or the inner three, because a plateau it will trust is two agreeing ratios and two ratios need three untouched whorls.

 Which arrangements carry a comb, and what each one reports. The largest comb mean in five arrangements at a rise of 0.005, all read by the same instrument at the same length, with the sampling band of 0.073 marked. Only the first is a placement rule; the other four are kinematic lattices with no rule in them, differing from one another only in how their azimuth errors are structured. Independent errors and errors with a memory leave nothing to read. A repeating error puts up a comb and names a partner that is not the lattice's. Errors inherited from the contact neighbours reproduce both the comb and the pair. Stems and cones

The comb was never the rule

A control is only as strong as the alternative it builds, and the earlier work built one that varied the rule while holding the disturbance fixed at independence. Five rounds of the angle-sequence thread, with what each claimed and what still stands — and why the next evidence has to come from an intervention rather than from a longer stem.

 Every rung has a window it can be read in. For each rung of the ladder, the range of lag windows in which the pair can be read: at least three times the smaller parastichy number, so the main comb has three teeth inside the window, and less than three times the larger, so the larger number cannot itself be a candidate spacing. The band is [15, 24) at 5/8, [24, 39) at 8/13, [39, 63) at 13/21. It is non-empty at every rung, because it is empty only when the larger number is no larger than the smaller. The line at 30 is the window the earlier work used everywhere: it sits inside the 8/13 band and outside the 13/21 one, which is what was mistaken for a ceiling. Stems and cones

The rung was not the instrument

The earlier work said the pair readout has a ceiling one rung above where it works, that this is arithmetic rather than statistics, and that no amount of stem fixes it. The arithmetic is right and gives a band of lag windows that is never empty; what was actually stopping the reading was an eight-node seed and a grid of 384 azimuths.

 How often three counters read a band's own pair, against the width of the band, on an ordinary stem. Bands of 40 to 320 organs slid up an ordinary stem grown at T = 300 over a rise from 0.05 to 0.0005, each read three ways. The counter without an index reads the pair at 65%, 66%, 87%, 88%, 88%, 88%, 89%, 100%, 100%, 92%, 54%, 23%, 8%, 2%, 1%, 0%, 0%, 0%; the counter families required to cross reads the pair at 65%, 66%, 87%, 88%, 88%, 88%, 89%, 100%, 100%, 92%, 54%, 22%, 8%, 2%, 1%, 0%, 0%, 0%; the counter with the index reads the pair at 100%, 100%, 100%, 100%, 100%, 100%, 100%, 100%, 100%, 100%, 100%, 100%, 100%, 100%, 100%, 100%, 100%, 100%. The index-free counter reads nine bands in ten or more only from 80 to 100 organs. The pattern itself

A counter that cannot be slid

The counter that needs no order of arrival follows each family into chains and counts them, and on an ideal lattice it agrees with the counter that does. On a stem whose rise falls it works only inside a band of widths, and outside the band it returns a pair rather than refusing: the rung below when the band is too narrow for the larger count, a pair on no rung when the band spans more than about a third of a rung of rise. The upper edge moves with the rate, so a width that is right on one stem is wrong on another, and on the fastest bijugate stem measured no width works at all.

 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. Where the angle comes from

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.

 How many scars a lineage carries for every growing point still alive. Deaths arrive at q times the standing count and the count grows by x a season, so the scars settle at q/(x − 1) — the curves. The dots are the ratio the expected counts actually reach after eighty seasons, and they agree to within 1.0 per cent. A bud waiting no delay at q = 0.1 carries 0.1250; a bud waiting one season at q = 0.1 carries 0.2192; a bud waiting two seasons at q = 0.1 carries 0.3135; a bud waiting three seasons at q = 0.1 carries 0.4128. Each curve runs to infinity at its own threshold, where the living stop outgrowing the dead. Branching and transport

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.

 What the round trip returns as the organs are displaced: golden angle, 300 organs. Thirty heads at each size of displacement, from none to two and a half spacings, each run through the round trip and sorted by what came back. The undisplaced head counts 21 and 34. At 0.25 spacings, 97% the undisplaced pair, 3% a neighbouring pair; at 0.5 spacings, 50% the undisplaced pair, 50% a neighbouring pair; at 0.75 spacings, 30% the undisplaced pair, 70% a neighbouring pair; at 1 spacing, 13% the undisplaced pair, 80% a neighbouring pair, 7% a straddling pair; at 1.25 spacings, 7% the undisplaced pair, 57% a neighbouring pair, 7% a straddling pair, 30% refused; at 1.5 spacings, 40% a neighbouring pair, 3% a straddling pair, 57% refused; at 2 spacings, 10% a neighbouring pair, 90% refused; at 2.5 spacings, 3% a neighbouring pair, 97% refused. 10 recovered intervals in all exclude the true angle. The pattern itself

A head displaced before it is counted

The round trip from a head's spiral counts back to its divergence angle was tested on heads whose every organ sat exactly where the rule put it. Displaced by a normal error of up to two and a half spacings, heads of 900 organs keep counting a pair from their own sequence and return intervals holding the true angle to a spacing and a half; heads of 300 organs move to the neighbouring pair by half a spacing and then refuse, nine in ten of them by two spacings. Every moved count brings in the family whose chord was third shortest. Of 898 heads recovered, 19 intervals miss the true angle and 17 of those by about a tenth of a degree — displacement makes the reading coarser and then silent, not confidently wrong.

 Every pair of death chances that reproduces each count, for one plant. A plant with a wait of two seasons, apices dying at 0.05 and buds at 0.15: a rate of 1.3351, 0.3098 scars per living point and 0.2225 of its scars left by apices. Each line is every pair of chances at that wait reproducing one count; the dashed lines either side are the same count one per cent high and low. The rate's and the scar share's lines cross at the plant's own pair, at an angle of 11.6°, so a one per cent error lets the pair slide along them — apex chances from 0.004 to 0.097 and bud chances from 0.100 to 0.198. The line for scars sorted by kind crosses the rate's at 51.5°, and read with it the same error leaves 0.044 to 0.056 and 0.134 to 0.166. Branching and transport

Two ways to die, three things to count

Giving a branching plant's waiting buds a death chance of their own leaves its counts a linear recurrence, but breaks the collapse onto the survival: the rate becomes the apex survival times the root of y^(d+1) = y^d + r^d, where r is the bud survival over the apex survival. The one-chance reading then names the wrong waiting time on 171 of 477 plants with waits of two to four seasons, shorter when the buds are the fragile ones and longer when the apices are. The two chances are separable from a rate and a scar share, exactly — but the two counts' loci cross at eight to sixteen degrees, so a one per cent error lets the chances wander by a factor of two. A third count is owed, and it is the scars sorted by kind.

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