Where the angle comes from

The stem that changed hands

A stem that never recovers from an ablation is supposed to end up somewhere worse than it started — a repeating block of angles, a pattern with the wrong counts in it. At the coarsest arrangement it ends up somewhere that is not worse at all: at 220.3125°, which is 360° minus the divergence it was cut from. The lattice is intact and its handedness is reversed.

Worth reading first: The organ that was taken away · Counting the spirals · The sequence has a memory.

Every arrangement of spirals has a hand. The organs of a stem step round the axis by a fixed angle, and that step is either clockwise or anticlockwise; a divergence of 139.6875° and a divergence of 220.3125° are the same lattice seen in a mirror. Every count is the same, every distance between organs is the same, every spiral is as tight — and a photograph of one cannot be superimposed on a photograph of the other.

Plants have hands too, and they are famously not fixed. A sunflower head is roughly as likely to run one way as the other, and both heads on one plant may differ. That is usually explained as a symmetry breaking that happens once, very early, and is then locked in. This essay is about a way of unlocking it after the fact — and the way is not subtle. Take two organs out of a settled stem and the rule, left to itself, sometimes rebuilds the whole pattern the other way round.

What was expected instead

The intervention has a well-established answer at finer arrangements. A stem cut in the middle of its front never returns to the divergence it had; it settles into an exactly repeating block of angles — five of them at the 5/8 arrangement, eight at 8/13 — precessing slowly around the axis, at an effective divergence far from anything the undisturbed rule produces.

The block a wrecked stem settles into is the count it was cut fromA stem is cut in the middle of its front and followed for 300 organs. It does not come back to its lattice; what it does instead is repeat a fixed sequence of divergences exactly, to the resolution of the azimuth grid. Each row shows that sequence twice over, one bar per organ, drawn to the same scale. At the 5/8 rung the sequence is five angles long and advances -36.1° per block; At the 8/13 rung the sequence is eight angles long and advances 22.5° per block. Every block is the smaller number of the pair that was cut, so the second attractor carries the first one's count — and every precession is one part in twice the block, which makes each of these a two-jugate arrangement with 10 and 16 rows.5/8 rungblock of 5-36.1° a blockand again208.8° on average8/13 rungblock of 8+22.5° a blockand again182.8° on average300 organs after the cut · 1536 azimuthsgenerated from a stated rule, not drawn to look right
Fig. 1 What a wrecked stem does at the two finer arrangements. The angles repeat exactly, in blocks whose length is one of the parastichy numbers of the lattice that was cut, and the effective divergence is over two hundred degrees.

Those orbits are unmistakable. Their motifs span a hundred and forty-eight and two hundred and twenty-six degrees; nobody looking at the sequence could think the stem had settled on a single angle. The prediction, extended downward, was that the coarsest arrangement would give a block of three.

What the coarsest arrangement does

It gives no block at all.

The wrecked stem is the lattice it was cut from, wound the other wayThe divergence of each organ placed after two were removed from a settled stem at a rise of 0.032, over the 220 organs following the cut. The upper line is the divergence the undisturbed stem holds, 139.6875°; the lower is 220.3125°, which is 360° minus it and therefore the same lattice with the opposite handedness. The sequence is thrown by the cut, wanders for a few dozen organs, and settles on the second line to four decimal places — 220.3125° against 220.3125° — where it stays. A counter shown the positions afterwards returns 3 and 5, the pair the stem was cut from. Nothing about the pattern has been lost; its chirality has been reversed, which at this rung a single removal cannot do.139.688°as grown220.313°its mirror060120180organs placed after the cutdivergencerise 0.032 · organs 3 and 4 back removed · counted 3/5generated from a stated rule, not drawn to look right
Fig. 2 The divergence of each organ placed after two were removed from a settled stem at a rise of 0.032. The upper line is what the undisturbed stem holds; the lower is 360° minus it. The sequence leaves the first and settles on the second.

The undisturbed stem settles at 139.6875°. The cut stem is thrown, wanders for a few dozen organs, and settles at 220.3125° — which is 360° − 139.6875° to the last digit that the arithmetic carries. It holds it for the remaining three hundred organs, with a scatter of a third of a degree at the azimuth grid used and a sixteenth of a degree at a grid three times finer.

Three of the four cells that fail to repair do this, at two different offsets and two different gaps. The agreement between them is exact rather than close: the same four decimal places, from cuts at different places in the stem’s history.

The wrecked stem is the lattice it was cut from, wound the other wayThe divergence of each organ placed after two were removed from a settled stem at a rise of 0.032, over the 220 organs following the cut. The upper line is the divergence the undisturbed stem holds, 139.6875°; the lower is 220.3125°, which is 360° minus it and therefore the same lattice with the opposite handedness. The sequence is thrown by the cut, wanders for a few dozen organs, and settles on the second line to four decimal places — 220.3125° against 220.3125° — where it stays. A counter shown the positions afterwards returns 3 and 5, the pair the stem was cut from. Nothing about the pattern has been lost; its chirality has been reversed, which at this rung a single removal cannot do.139.688°as grown220.313°its mirror060120180organs placed after the cutdivergencerise 0.032 · organs 2 and 3 back removed · counted 3/5generated from a stated rule, not drawn to look right
Fig. 3 The same measurement after a different pair of organs is removed. A different intervention, the same destination, to four decimal places — which is what makes this an attractor of the rule rather than a coincidence of one run.

The counts come back with it

A divergence that happens to sum to 360° with another one would be a curiosity if nothing else agreed. Everything else agrees.

A counter shown the wrecked stem’s positions, with no access to the angle it was built from, returns 3 and 5 — the pair the stem was cut from. The three shortest hops are five, three and eight organs, in that order, exactly as before. The spacing between neighbours is the same, because it is a function of the same lattice.

Which offsets give short hops, at a rise of 0.032The two lowest points are at 3 and 5, and those are the parastichy numbers. Offset 1 is high because a hop of one node is at least the rise, which is what makes a stem easier to count than a disc.00.5001102030index offsetmedian hop between node i and node i+m35300 nodes, 34 offsets triedshortest at 3 and 5
Fig. 4 The hop lengths of the arrangement before the cut. Five organs is the shortest step, three the next, eight the next. After the cut the same table comes out of the mirrored stem, in the same order.
A stem unrolled: 70 nodes at 139.69° with a rise of 0.032 circumferencesThe counter is shown these coordinates and the circumference, and finds 3 parastichies one way and 5 the other. The faint strips left and right are the same stem: a family leaving one edge re-enters at the other.3 and 5rise 0.032 · divergence 139.69°counted 3 and 5, opposed
Fig. 5 The lattice as grown, unrolled. The two contact families run three one way and five the other, and the counter is shown these coordinates without being told the angle.
A stem unrolled: 70 nodes at 220.31° with a rise of 0.032 circumferencesThe counter is shown these coordinates and the circumference, and finds 3 parastichies one way and 5 the other. The faint strips left and right are the same stem: a family leaving one edge re-enters at the other.3 and 5rise 0.032 · divergence 220.31°counted 3 and 5, opposed
Fig. 6 The lattice the wrecked stem settles into, drawn at the divergence it actually reaches. Every count and every hop length is what the previous figure has; the families lean the other way.

So the stem has not been damaged into some third thing. It has been moved from one member of a pair of equivalent lattices to the other, and there was never a sense in which it was on the first one rather than the second except that it had been started there.

The arithmetic of why a mirror is the same lattice

The identity is worth writing out, because it is what makes the result a statement about handedness rather than about a coincidence between two numbers.

An organ i on a cylindrical lattice sits at azimuth iδ and at height ih, with the azimuth read around a circle of circumference one. The distance from organ i to organ i + k depends on δ only through the wrapped quantity kδ modulo a whole turn, and it depends on it only through the magnitude of that wrap, since a step of +x and a step of −x are the same distance either way round the cylinder. Replace δ by 360° − δ and every wrap changes sign and keeps its size. So every hop length is unchanged, for every k, exactly.

Everything a counter uses is a hop length. The parastichy numbers are the two offsets with the shortest hops; the incoming number is the shortest hop longer than those; the whole ladder of transitions is solved from the same table. None of them can distinguish a lattice from its mirror, and none of them should: the two point sets are congruent, and a rule that placed organs by repulsion alone cannot prefer either.

What does distinguish them is the sign of the step, which is not a distance and is not in the table. It is in the sequence of azimuths and nowhere else — which is to say, it is in the order the organs were made in. That is the sense in which handedness is a fact about growth rather than about form, and it is why a measurement made on a photograph reports it and a measurement made on a neighbour graph cannot.

What the transient looks like

The stem does not jump. It is thrown by the cut — the organ placed immediately afterwards is sixty-two degrees from where it would have been — and then spends somewhere between thirty and sixty organs producing divergences that belong to neither lattice, before the sequence tightens onto the mirrored value and stays there.

That transient is not noise and it is not a slow drift. Watched closely it is the rule doing what it always does: each organ placed at the least of a profile, with the profile still deformed by the two vacancies and by every organ placed since. The vacancies pass out of the neighbourhood after a few dozen placements, and from then on the rule is being asked its ordinary question about an arrangement that is already nearly the mirror.

The length of the transient is the reason the recovery test needs a hold. A test that asked only whether the last few angles are close to a settled value would call a wandering stem settled somewhere in the middle of this stretch. The test used here requires every divergence from some organ onward to stay within a degree and a half of the value being tested, with at least sixty of them to check, which is why a wrecked stem is reported as wrecked rather than as slowly recovering.

A single organ cannot do it

This is the part that makes the result an experiment rather than an observation about symmetry. At this arrangement, every single-organ removal repairs itself — all five offsets of the front, at three cut points four dozen organs apart, back on 139.6875° within thirty-nine organs.

The next organ moves for the last 5, and for no othersOne row per organ removed, counted back from the tip of a stem at a rise of 0.032 whose counted pair is 3 and 5. Removing any of the last 5 moves the next organ by 8.7° to 149.1°; removing an older one moves it by at most 1.41°, which is under the azimuth grid. The boundary is at 5, and 5 is the larger parastichy number — so the experiment counts the spirals without measuring an angle.organ removed, counted back from the tiphow far the next organ moves, in degrees1139.7°279.0°342.7°4149.1°58.7°— the front ends here60.9°71.2°81.4°90.2°100.7°110.5°120.2°130.0°140.0°150.2°160.0°rise 0.032 · pair 3/5generated from a stated rule, not drawn to look right
Fig. 7 Removing one organ at a time from the coarse stem. The displacements are large — up to a hundred and fifty degrees — and every one of them is repaired. Nothing at this arrangement is thrown far enough to leave the basin it started in.
A cut four back is undone after 38 organsThe divergences of a stem whose organ four places back was removed, against the same stem uncut. The sequence is thrown by 149° and is back within 1.5° of its settled 139.7° after 38 organs, and stays there for the remaining 262. This is the rule correcting itself: an organ placed to one side of its minimum leaves a gap that pulls the next one back.100150200250300050100organs placed after the removaldivergence, in degreesback on the latticerise 0.032 · cut 4 backgenerated from a stated rule, not drawn to look right
Fig. 8 The offset that is beyond repair one arrangement finer, at this one. The sequence is thrown by a hundred and forty-nine degrees, wanders, and comes back to where it was.

Two organs can do it and one cannot, at four of sixty-four pairs of offsets. That gives the pair of lattices a quantitative separation: they are further apart than one removal throws a stem and closer than two removals at the right offsets. It is a statement about the size of a basin, made with an intervention that has a size.

A second cut moves the next organ, and does not move the boundaryEvery pair of organs that can be taken out of a settled stem at a rise of 0.032, where the pattern is 3/5. A row is the offset of the nearer organ removed, counted back from the tip; a column is how many further places back the second one sits. The shade is how far the next organ ends up from where it would have been, from under a degree in the palest cells to a half-turn in the darkest. The run of shaded rows ends at 5, which is the larger parastichy number, and every row below it is blank across the whole width — the largest displacement anywhere past the front is 2.34°. So the nearer of the two organs decides whether the removal is felt at all, and the second one, wherever it is put, cannot make the pattern notice an organ it was not going to notice.816117891311411391411403811542808477817779441950414244414442915014714915014914914914981079989880210111110121111112121221210000000005 = 5123456789123456789nearer organ,places backgap to the second organ, in placesdisplacement of the next organ, in degrees · pair 3/5rise 0.032 · 400 organs before the cutgenerated from a stated rule, not drawn to look right
Fig. 9 The whole two-organ experiment at this arrangement. The four cells that never repair sit at offsets two and three with small gaps — inside the front, where the two vacancies are close enough for the repair of one to spoil the repair of the other.

The fourth cell is not a mirror, and it is worth keeping

Of the four cells that fail, three settle on the mirror and one does not. A cut at offsets two and four settles at 258.73°, whose three shortest hops are four, three and seven — a 3/4 lattice, which is neither the original nor its mirror.

It is reported rather than smoothed away because a result that held in four cells of four would be a weaker claim than one that holds in three: the fourth cell is what says the mirror is a destination the rule can reach rather than the only thing available once a coarse stem is disturbed. There are at least two places to go, and most of the failures go to the same one.

The 3/4 lattice is also interesting on its own account. Four is not a Fibonacci number and 3/4 is not a rung of the ladder a golden-angle stem walks; it is a lattice the rule can sit on at this rise without being on the ladder at all. Its divergence, 258.73°, has a mirror at 101.27°, which nothing here reaches — so the cell that does not go to a mirror does not go to a mirror in both senses. What it shares with the other three is that it is a lattice: a single divergence held for three hundred organs, not a block of angles, and a counter shown its positions returns a pair.

Taken together the four cells say the coarse arrangement has at least three resting places within reach of a two-organ disturbance, and that all of them are lattices. That is a different situation from the finer arrangements, where every failure lands on an orbit, and it is the shape of the difference this essay is about.

What it would mean on a plant

Handedness is one of the four things this collection’s survey specification asks a real census to record, and it is the one nobody reports. Spiral counts are published constantly; the counting radius, the element count, the rotational symmetry and the hand are the fields that turn a count into a measurement, and they are usually treated as context.

The prediction here gives the hand something to do. On a coarsely patterned shoot — three and five, which is common on young stems and on many rosettes — removing two nearby primordia should, at a minority of offsets, produce a shoot whose subsequent phyllotaxis runs the other way, with the same counts as before. That is a large, obvious, photographable outcome from a small intervention, and it is one that no account in which the hand is fixed early and locked in can produce.

It is also a test with an unusually good null. If the hand were fixed by something upstream of the placement rule, an ablation could not change it at any offset; if the hand is nothing but which of two equivalent lattices the neighbourhood currently sits on, it should be changeable by a disturbance of about the size measured here. Those two predictions differ in kind rather than in degree.

Every transition as the rise fallsThe pair climbs 1/2 → 2/3 → 3/5 → 5/8 → 8/13. Consecutive transitions are 0.381, 0.382, 0.382 of the previous rise — 1/φ² is 0.3820.0.4000.6000.8001-2-1.50-1log₁₀ of the rise between nodes (falling to the right is the plant growing)log₁₀ of the larger parastichy number2/33/55/88/13500 rises, shortest vectors recomputed at eachratio 0.3819 against 1/φ² = 0.3820
Fig. 10 Where the coarse arrangement sits on the ladder. The pair a cylindrical lattice carries is set by the rise, so an experiment can be pointed at the arrangement it needs by choosing a shoot at the right stage rather than by hoping.

Why the coarse arrangement and not the fine ones

The finer arrangements do not do this, and the reason is visible in what they do instead. At 5/8, twenty-two of thirty-six pairs of offsets fail to repair, and every one of them lands on a repeating block rather than on a lattice. The mirrored divergence, 223.22°, is not among the values reached.

The natural reading is about how much room there is. A coarse lattice has five organs in its front and two vacancies in it are a large fraction of the neighbourhood; the rule is thrown a long way, and the two nearest places to land are the two hands of the same lattice. A fine lattice has thirteen organs in its front, two vacancies are a small fraction of it, and what the rule finds nearby is a rearrangement rather than a reflection.

That is a reading rather than a measurement, and the measurement that would test it is available: the same table at 5/8 with three organs removed, which is a larger disturbance on a bigger neighbourhood. Nothing here has run it.

The block is one of the two numbers, and not always the smallerEvery lattice a single organ was removed from, one row each: golden, rise 0.020 carrying 3/5; golden, rise 0.013 carrying 5/8; golden, rise 0.008 carrying 5/8; golden, rise 0.005 carrying 8/13; Lucas, rise 0.020 carrying 4/7; Lucas, rise 0.013 carrying 4/7. The two open ticks on each line are that lattice's own parastichy numbers; the filled dots are the blocks the stems that never recovered settled into, one per offset that failed. The claim this table was built to test is that the block is the smaller of the two, which held at the two rises it was first measured at. It does not hold here: 3/5 gives 5, 5/8 gives 5 and 8, 4/7 gives 4 and 7. What survives is weaker and still worth something — every filled dot but 1 sits on one of that row's own ticks, so the orbit carries a count of the lattice it was cut from, and which of the two it carries is decided by the offset rather than by the pattern.13579111315golden, rise 0.020pair 3/5golden, rise 0.013pair 5/8golden, rise 0.008pair 5/8golden, rise 0.005pair 8/13Lucas, rise 0.020pair 4/7Lucas, rise 0.013pair 4/7block, in organs — open ticks are the lattice's own pairfilled dark where the block is the larger of the pairone organ removed · 400 organs before the cutgenerated from a stated rule, not drawn to look right
Fig. 11 The blocks a wrecked stem settles into across several arrangements and both branches. Every filled dot is an orbit; the coarse arrangement contributes nothing to this table, because what it settles into is a lattice rather than an orbit.

What this does not say

It does not say the rule prefers one hand. It has no preference: the rule is exactly symmetric under reflection, and a stem seeded at 220.3125° stays there as firmly as one seeded at 139.6875°. What the measurement shows is that the barrier between them is finite and has a size that an intervention can be compared with.

It does not say a plant’s handedness is unfixed. It says that in a model where the hand is nothing but a property of the current arrangement, a two-organ ablation flips it at some offsets. Whether a meristem carries anything else that remembers the hand is exactly what the experiment would find out, and the model cannot answer it.

It does not say the mirror is reached from every failure. Three of four cells at one arrangement, and none at all one arrangement finer. This is a property of coarse patterns and it is stated as one.

And it does not say the sequence is noiseless. The wrecked stem’s divergence scatters by a third of a degree at the grid the table is computed on. That scatter is the grid — at three times the resolution it falls to a sixteenth of a degree while the mean does not move — which is the same discretisation that produced a spurious repeating block in an earlier reading of this run, and which has its own essay.

The check that would refuse it

The claim is three things at once, and all three are asserted while the figures are drawn.

The settled divergence of the wrecked stem has to be within a twentieth of a degree of 360° minus the divergence it was cut from. That tolerance is a fifth of the azimuth grid’s own step, and the three cells that satisfy it satisfy it to 0.0000°, so nothing is near the threshold. A run that landed near the mirror rather than on it would be reported as near.

A counter shown the wrecked stem’s positions has to return the pair the stem was cut from. This is the strong half: two angles summing to 360° is arithmetic, and a counter agreeing about both parastichy numbers is a statement about the point set.

And not every failing cell may be the mirror. If all four went to the same place the figure would be consistent with the rule having exactly one destination after a coarse stem is disturbed, which is a different and less interesting claim; the check requires at least one cell that does something else, and the 3/4 lattice at 258.73° is what supplies it.

Shares its objects with

Essays that name at least two of the same things, and that neither author linked.

  • A stem on the other branch — both name ablation, attractor, counting blind, honest limits, initial condition, lattice, measurement, parastichy pair, the placement rule, rise, rung
  • A front with no middle — both name ablation, divergence angle, equilibrium, honest limits, lattice, measurement, parastichy pair, the placement rule, rise, rung
  • The pattern the cut leaves behind — both name ablation, attractor, counting blind, divergence angle, equilibrium, honest limits, lattice, measurement, parastichy pair, the placement rule
  • Two accounts of one number — both name ablation, attractor, counting blind, honest limits, lattice, measurement, parastichy pair, the placement rule, rise, rung
  • The response with a hole in it — both name ablation, counting blind, divergence angle, honest limits, measurement, parastichy pair, the placement rule, rise, rung
  • Two-ranked, by two different routes — both name ablation, attractor, divergence angle, equilibrium, honest limits, initial condition, measurement, the placement rule, rise

Named objects

A flat tag is an object no other essay names yet.

AblationAttractorBasinCounting blindDivergence angleEquilibriumHandednessHonest limitsInitial conditionLatticeMeasurementParastichy pairThe placement ruleRiseRung