Phyllotaxis
the spirals are counted, not admired
Concept

Round trip — where it appears

Building a pattern from a stated number, then recovering that number from the pattern by a route that never sees it. It is the strongest check available without data, and this collection uses it on divergence angles, growth factors and rises.

Named by 18 essays across 4 fields — each of them below, with the objects they name alongside it.

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 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 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 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.

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.

lattices · Counting Two combs, at a rise of 0.005. The autocorrelation of 760 divergence angles from one stem held at a rise of 0.005. The filled teeth are the lags at multiples of 8; the open teeth are the second comb, at the same spacing offset by 5. Reading the spacing off the first and the offset off the second gives the pair 8 and 13, which is what the position counter reports for the same stem — from angles alone, with no coordinate anywhere in the calculation.

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 The ratio of two whorls' line counts is the growth factor raised to the clock's own power. Each curve is W^p for one law: flat for a clock that advances the angle at a constant rate, W for one that adds a constant length at the opening, W² for a constant area and W³ for a constant volume. At 3.20× per turn those are 1.00, 3.20, 10.24, 32.77. So a count of lines in two successive whorls, divided, and read back through a growth factor the curve already gives, names the law — and the four are further apart the faster the shell expands.

What the growth lines carry

A shell's curve says nothing about how fast the animal grew, and its growth lines say all of it. Under a law that holds the pth power of the radius constant per unit time, the time spent crossing one whorl is proportional to the change in that power across it — so the lines in successive whorls stand in the ratio of the growth factor raised to p, and each whorl holds the count the closed form predicts to within the one line rounding can move. Dividing two counts and taking the logarithm against a growth factor the curve already gives returns p: 17 of 20 readings name their own law, the furthest 0.012 from a whole number. The other three are not wrong, they are uncountable — at 4.5 per turn under a volume clock the inner whorl of the pair holds one line.

shells · Spiral 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.

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.

shells · Spiral 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.

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.

cylinder · Noise transport A shell section square on and seen 30 degrees off. The same three turns of a spiral built at 3.2 per turn, drawn as a camera normal to the coiling plane sees it and as one 30 degrees away from normal does. The tilted view is the plane compressed by 0.8660 along one direction. A fit to the first returns 3.200000 with a residual of 1.8e-15; a fit to the second returns 3.19527 with a residual of 0.07763. Nothing in the second picture says it is not a shell.

A section seen from the wrong angle

A photograph of a shell section taken off the normal is the coiling plane compressed along one direction by the cosine of the angle, and nothing in the picture says so. The fit that recovers a growth factor is moved by it — half a turn seen twenty degrees off gives a band of answers 23.7 per cent wide as the span's starting point moves round the shell, centred almost exactly on the right answer, so it is a spread and not a bias. The caliper measure is exactly immune at every tilt and every aim, because a projection scales all three points on a line through the centre by the same factor. And the fit's residual names the tilt to three decimal places, which makes this the rare error a section reports about itself.

shells · Spiral fit 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.

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 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.

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.

lattices · Recovery A head of 300 organs as the rule places it, and twisted by 2 rad. Two heads at the golden angle, 300 organs each. On the left every organ is where Vogel's rule puts it, and the round trip counts 21 and 34 and recovers 137.515°. On the right the same organs turned about the centre by 2 radians times their radius over the head's, so the rim turns by 2 radians and the centre not at all; the round trip counts 34 and 47, recovers 137.80°, missing, with an interval of 0.230° that excludes the true angle.

A twist is a divergence

Recovering a head's divergence angle from its spiral counts survived independent displacements of whole spacings, moving to a neighbouring pair and then refusing rather than misleading. Displacements with a direction are harder on it in only one case. A head pressed to an aspect ratio of 2.25, spread at the rim by eighty per cent or sheared with a slope of 1.6 is still counted as its own pair or its neighbour, and recovered inside its interval. A head twisted — each organ turned about the centre in proportion to its radius — is not: past a turn of the rim of about a radian and a half the counts leave their sequence and the recovered angle misses, by up to sixty-one degrees, because a twist changes the angle between one organ and the next. The round trip is not fooled. It is reporting the angle the twisted head has.

lattices · Recovery A 900-organ head twisted by 0.5 radians at the rim, with the bands its spirals are counted in. Every organ turned about the centre by 0.5 times its radius over the head's. The rings mark the inner annulus, 0.35 to 0.6 of the radius, and the outer, 0.7 to 0.95. The inner annulus counts 34 and 55; the outer 34 and 89 — skips one; the single band, 0.55 to 0.95, 34 and 89 — skips one. A count that is not two consecutive Fibonacci numbers flags the twist; on this head that happens from half a radian, and the single band is misled only from six.

The count sees the twist first

A twisted head recovers a changed divergence, and the check proposed for it was two annuli: the twist's extra angle falls with radius, so an inner and an outer annulus should disagree. Read on golden heads of 900, 2,400 and 9,000 organs, they never do in time. Their intervals separate at eight radians on 900 organs and never on the larger heads, always after the ordinary reading has been misled — from six radians on 900 organs and from two on 2,400 and 9,000. What catches the twist first, at every size and on every seed, is the count: at half a radian to three quarters some band stops returning two consecutive Fibonacci numbers — 34 and 89, 89 and 233 — which no untwisted golden head, clean or displaced, ever does.

lattices · Recovery How much a twist of the rim changes the divergence between neighbouring organs, across a 900-organ head, for six shapes of twist. A twist of 0.25 radians at the rim, shaped as a·(r/R)^p, changes the angle between one organ and the next by a·(p/2)·ρ^(p−2)/(N − 1) at a distance ρ from the centre: the six lines are exponents 0.5, 1, 1.5, 2, 3, 4. At two the change is the same everywhere — 0.0159° — which is a different divergence angle; below two it is largest at the centre, above two at the rim. The shaded ring is the outer annulus, and the band between dotted lines is the flag's resolution on an untwisted 900-organ head: grown 0.0125° off the golden angle it counts consecutive Fibonacci numbers, 0.015° off it does not. At this twist the outer annulus passes it for exponents 2, 3, 4.

The flag reads the angle, not the twist

A twisted seed head is caught first by its counts: at half a radian some band stops returning two consecutive Fibonacci numbers, which an untwisted golden head never does. Twist the head in proportion to the square of the radius instead and the organs land exactly where a head grown at the golden angle plus a/(N − 1) puts them — to a billionth of a spacing — so the two read the same pairs in every band, and the flag fires on both. The flag detects a divergence that is not golden, and it has a resolution: 0.015° on 900 organs, 0.01° on 2,400, 0.003° on 9,000. Every twist shape from a half to four is flagged exactly when its change to the divergence in the outer annulus passes that resolution. So a flagged head is not golden, and nothing in its counts says whether it was twisted or grown that way.

lattices · Recovery 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.

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.

branching · Lsystem Four hundred plants whose seasons are sometimes bad, each measured against the count the average season predicts. A two-season wait, and every season bad with chance 0.1, killing each growing point with chance 0.5; good seasons kill with 0.0556, so the chance averaged over seasons is 0.1. Four hundred sequences of forty seasons, each plant's living count divided by the count the independent model expects at 0.1. The dashed line at one is the expectation over every sequence of seasons; the mean of these four hundred is 1.073 at season forty, as near as a sample carried by its luckiest few plants comes. The solid line is the median plant, 0.540 of the expected count by season forty, and the band holds the middle ninety per cent, from 0.080 to 3.640.

A bad year does not average out

Let every growing point on a plant share one season's death chance, bad one season in ten and good otherwise, with the average held at 0.1. Averaged over every sequence of seasons the counts are exactly the independent model's. But no plant is an average over sequences: with bad years at 0.5 a plant settles on a rate of 1.3001 against the expected 1.3190, the median plant holds 54 per cent of the expected count by season forty, and the scar share never settles, because it is set by how many seasons ago the last bad year was — 1.35 the season after one, 0.17 twelve seasons on. So the reading of a plant's waiting time from a rate and a scar share gets worse the longer it runs: right for 57 per cent of plants over ten seasons, 22 per cent over eighty.

branching · Lsystem One plant's scars dated by season, and the wait they name when the recurrence is run through its own seasons. A plant with a two-season wait, grown point by point for 20 seasons from one apex; each season is bad with chance 0.1, when each point dies with chance 0.5. Top: the share of living points that died each season, read from that season's scars, with the bad seasons marked. Bottom: the living count on a logarithmic scale, and the counts a wait of one, two and three seasons predicts when the recurrence is run from one apex through these same seasons' death shares. The two-season line follows the plant; dated, the scars name a wait of 2. Read as two totals — the rate after the first 5 seasons and scars over living points — the same plant names 1.

Scars with dates on them

A bad season shared by every growing point wrecks the reading of a branching plant by two totals: over eighty seasons the rate and the scar share name a two-season wait for 22 per cent of plants. Date the scars — by position along a shoot, by growth ring — and each season's death chance is read off its own scars, so the bad years stop being noise and become a known input. Running the branching recurrence through the plant's own seasons names the wait for 94 per cent of plants over twenty seasons whether bad years kill a tenth of the points or seven tenths, and for every plant over eighty. What a plant cannot read from its own scars is the climate: the averaged chance comes only as fast as seasons do.

branching · Lsystem

Named alongside it

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

Honest limitsFibonacciClaim testingIdentifiabilityMeasurementDivergence angleMeasurement errorParastichyRefusalCylinderInterval estimateParastichy pair

All concepts