Stems and cones

What a cut costs a whorl

A bijugate pattern is an ordinary lattice seen twice over, so the account that says a wrecked stem's repeating block is the repeat unit of the lattice underneath has a specific prediction here: three and five. It gets six and ten. And the thing a single missing organ does destroy on a whorled stem is the one property its counts cannot see.

Worth reading first: Counting the spirals · The organ that was taken away.

A bijugate pattern puts two organs at every node instead of one, half a turn apart, and steps between nodes by a divergence of its own. Its parastichy numbers are both even; a stem counted at four and six is a stem whose two families come in pairs. The arithmetic of it is settled here already: a k-jugate lattice is the ordinary lattice at k times the divergence and k times the rise, seen k times over, with every count multiplied by k, and the bijugate limit divergence is exactly half the golden angle.

That makes it the sharpest available test of one particular account of what a stem does after an ablation it cannot repair. The account says the repeating block it settles into is a repeat unit — the count of the ordinary lattice underneath a jugate arrangement. On a bijugate stem there really is an ordinary lattice underneath, and its counts are half the bijugate ones. So the prediction is unambiguous, and it is different from every other account’s.

The rule has to be extended, and only in one place

The placement rule this collection uses puts one organ at a time at the least of a repulsion profile. It cannot make a whorled pattern: nothing in it produces two organs at one height.

So the rule is extended in the smallest way that does. Organs arrive k at a time at one height, and each one is placed at the least of the same profile against everything already there — its own whorl-mates included. The exponent, the neighbourhood, the window and the argmin are untouched, and the k-fold symmetry is deliberately not imposed: the second organ of a whorl is placed by the same argmin as the first, and where it lands is a result.

three stems: 1, 2, 3 primordia at a time. 1-jugate at 137.51° counts 2 and 3; 2-jugate at 68.75° counts 2 and 4; 3-jugate at 45.84° counts 3 and 6. Every count shares the factor k, and dividing it out leaves an ordinary lattice: what a k-jugate stem is, exactly, is k copies of an ordinary one wrapped k times round.
Fig. 1 What a whorled arrangement looks like, at three jugacies, drawn from the geometry rather than grown. Two organs at a node, three at a node, and the ordinary one for comparison.

Grown that way at three rises, the rule produces lattices with rotational symmetry of order 2, measured on the positions afterwards by rotating the whole point set and requiring every rotated organ to land on an existing one. Their pairs are 4/6 at rises of 0.026 and 0.013 and 6/10 at 0.0065 — which are rungs of the bijugate ladder this collection derived from the arithmetic, 2/4, 4/6, 6/10, 10/16.

A bijugate stem answers in pairs, once the half turn is taken out. Removing one organ from a stem grown by a rule that places three at a time, at a rise of 0.0065, where the pattern counts 6 and 9 and has rotational symmetry of order 3. The open circles are the displacement of the next organ as it comes out of the arithmetic, which reaches 120° at offsets where the stem has demonstrably not been disturbed — the two organs of a whorl are interchangeable, so calling the other one "next" is a relabelling and not a movement. The filled circles are the same numbers read modulo 120°, which is the only way a 3-jugate divergence is defined. Read that way the response is a run of equal pairs — 47.6°, 47.6°, 47.6°, 22.3°, 22.3°, 22.3° — ending at 9, the larger parastichy number, with everything past it under 0.2°. Two organs of one whorl give the same answer to the last digit, which is the pattern's symmetry showing up in an experiment.
Fig. 2 A three-jugate stem at the fine rise, cut one organ at a time. What a removal costs is read the same way at every jugacy.

Counting a pattern with no genetic spiral needs a counter that never sees an index: the short repeated displacement vectors are recovered from the positions, each is traced into chains, and the chains are counted, which is what a thumb on a pine cone does and what an index-based counter cannot.

The first thing the experiment changes is the measurement

Removing one organ from a whorled stem breaks the symmetry, and the displacement of the next organ has to be read differently because of it.

The two organs of a whorl are interchangeable. An intervention that swaps which one the rule calls “next” reports a displacement of half a turn while nothing has moved. Taken at face value the response never ends: at the 6/10 arrangement, offsets eleven and twelve report displacements of 0.5° and 179.5°, and both stems demonstrably recover within a handful of organs.

A bijugate stem answers in pairs, once the half turn is taken out. Removing one organ from a stem grown by a rule that places two at a time, at a rise of 0.0065, where the pattern counts 6 and 10 and has rotational symmetry of order 2. The open circles are the displacement of the next organ as it comes out of the arithmetic, which reaches 180° at offsets where the stem has demonstrably not been disturbed — the two organs of a whorl are interchangeable, so calling the other one "next" is a relabelling and not a movement. The filled circles are the same numbers read modulo 180°, which is the only way a 2-jugate divergence is defined. Read that way the response is a run of equal pairs — 68.0°, 68.0°, 42.9°, 42.9° — ending at 10, the larger parastichy number, with everything past it under 0.5°. Two organs of one whorl give the same answer to the last digit, which is the pattern's symmetry showing up in an experiment.
Fig. 3 Removing one organ from a bijugate stem, offset by offset. The open circles are the displacement as it comes out of the arithmetic and the filled ones are the same numbers read modulo half a turn, which is the only way a bijugate divergence is defined.

Read modulo 180°, the response is a clean run that ends at ten, which is the larger parastichy number, with everything past it under half a degree. The boundary result therefore survives multijugacy — and it needed to be checked, because a rule that gives the count on ordinary stems and half a turn of noise on whorled ones would be a rule with a hidden assumption in it.

And the response comes in pairs

The clean part. Removing the organ one place back and the organ two places back displaces the next organ by the same amount to the last digit — 67.97° at both — then three and four by 42.89° at both, five and six by 20.63°, seven and eight by 76.64°.

That is the pattern’s own symmetry appearing in an experiment rather than in a picture. The organ one place back and the organ two places back are the two members of one whorl, and they are equivalent under the rotation that maps the lattice onto itself, so the rule cannot distinguish them. It is the cheapest test on this page that a stem really is bijugate, and it costs one extra ablation.

It is also a test a photograph cannot do. A pattern can look whorled and be an ordinary lattice whose two families happen to share a factor; this collection has built exactly that case, an ordinary lattice at 180.5° counting two and four with rotational symmetry of order one. Counts cannot separate the two. A response constant on whorls can.

How a whorled stem’s divergence is even defined

There is a measurement question here that the ordinary case does not raise, and getting it wrong produces a result rather than an error, so it is worth setting out.

On an ordinary stem the divergence is the angle from each organ to the next, and a settled stem gives the same number over and over. On a bijugate stem the angle from each organ to the next alternates: half a turn to its whorl-mate, then whatever is left to the next whorl’s first organ. A sequence measured that way has no settled value at all, and a first version of this work duly reported an undisturbed bijugate stem as a period-two orbit — a perfect lattice reported as already wrecked.

The quantity that behaves is the whorl divergence: the angle from organ i to organ i + k, which is the same rank one whorl up. On a settled bijugate stem at the finest rise here it is 68.98° with a scatter of 0.11°, which is within a quarter of a degree of half the golden angle. It is reduced modulo one k-th of a turn for the same reason the displacement is, and for a reason this collection already had: a k-jugate divergence is only defined modulo 360/k, because adding that much leaves the point set identical.

So a whorled stem needs two conventions an ordinary stem does not, and both of them are forced rather than chosen. Every recovery time and every block below is measured on the whorl divergence, folded.

What the block is, and what it is not

At the 6/10 arrangement, eight of thirteen offsets fail to repair, and their orbits are wide — motifs spanning a hundred and seven to a hundred and fourteen degrees, so they are orbits and not the azimuth grid rounding a constant. The blocks are six and ten.

Six and ten are the two parastichy numbers of the bijugate lattice. The ordinary lattice it is two copies of has the pair 3/5, and neither three nor five appears at any offset. The repeat-unit account predicted exactly those two numbers, on the strongest ground it had, and they are absent.

The trijugate arrangement is drawn at the fine rise and nowhere else, and that is a refusal rather than an omission. At 0.013 and 0.026 the three-organ response does not run out to its larger parastichy number — the pair is 3/6 and the front stops at three — so the boundary result that holds at every bijugate rise, and at the fine trijugate one, does not hold there. Whether that belongs to the jugacy or to the rise is one sweep away and is not settled here.

The two coarser bijugate rises do not produce orbits at all, and that is reported rather than folded in. At 0.026 the wrecked stem takes a constant whorl divergence — 23.44° at two offsets and 156.56° at two others, with a motif spanning 0.00° — and at 0.013 the motifs span two to five degrees, which is ten to twenty steps of the azimuth grid and not an orbit either. A period read off either would be the grid’s period rather than the pattern’s, which is a mistake this collection has already made once and now tests for.

A bijugate stem answers in pairs, once the half turn is taken out. Removing one organ from a stem grown by a rule that places two at a time, at a rise of 0.013, where the pattern counts 4 and 6 and has rotational symmetry of order 2. The open circles are the displacement of the next organ as it comes out of the arithmetic, which reaches 180° at offsets where the stem has demonstrably not been disturbed — the two organs of a whorl are interchangeable, so calling the other one "next" is a relabelling and not a movement. The filled circles are the same numbers read modulo 180°, which is the only way a 2-jugate divergence is defined. Read that way the response is a run of equal pairs — 71.2°, 71.3°, 34.2°, 34.2° — ending at 6, the larger parastichy number, with everything past it under 1.2°. Two organs of one whorl give the same answer to the last digit, which is the pattern's symmetry showing up in an experiment.
Fig. 4 The middle arrangement. The response is the same shape — pairs, a boundary at the larger count, nothing past it — and what the unrepaired offsets settle into is not an orbit, so no block is reported from this row.

The symmetry does not come back

The result that has nothing to do with blocks, and is the one an experimenter could see fastest.

Every wrecked bijugate stem here has rotational symmetry of order 1 where it had 2, at every rise, and none of them recovers it in three hundred organs. The pattern goes on producing a lattice — a counter shown the positions returns a first parastichy number that makes sense — and the property that made it bijugate is gone permanently.

That is worth putting beside what this collection already says about jugacy: the counts cannot decide it. An ordinary lattice and a genuine bijugate one can carry the same pair, and only the symmetry of the point set separates them — which is the inference a wrecked spiral stem breaks from the other direction. So jugacy is the one property of a whorled pattern that a count cannot see, and it is the property a single missing organ destroys.

An experimenter looking for the cheapest observable in this whole thread should probably look here. Counting parastichies on a shoot is work; noticing that a shoot which used to have two organs per node no longer does is not.

A bijugate stem answers in pairs, once the half turn is taken out. Removing one organ from a stem grown by a rule that places two at a time, at a rise of 0.026, where the pattern counts 4 and 6 and has rotational symmetry of order 2. The open circles are the displacement of the next organ as it comes out of the arithmetic, which reaches 179° at offsets where the stem has demonstrably not been disturbed — the two organs of a whorl are interchangeable, so calling the other one "next" is a relabelling and not a movement. The filled circles are the same numbers read modulo 180°, which is the only way a 2-jugate divergence is defined. Read that way the response is a run of equal pairs — 63.8°, 63.7°, 33.3°, 33.3° — ending at 6, the larger parastichy number, with everything past it under 0.9°. Two organs of one whorl give the same answer to the last digit, which is the pattern's symmetry showing up in an experiment.
Fig. 5 The coarsest of the three. The pairing is exact here too, the boundary is again the larger count, and what the unrepaired offsets settle into is a single whorl divergence held constant — a motif spanning nothing at all.

How often a single loss would cost a plant its jugacy

The symmetry result has a rate attached to it in the same measurement, and putting the two together turns an observation into a prediction about populations.

Eight of the thirteen offsets tried at the 6/10 arrangement never repair, and every wrecked stem loses its symmetry permanently. So a single organ lost at a randomly chosen recent position destroys jugacy about three times in five, and the remaining two in five are repairs that leave the pattern as it was.

That is a large number for an event that is not rare. A shoot loses organs — to insects, to frost, to a passing animal — and it loses them from the youngest tissue, which is exactly the region the offsets in this census cover. Under the model a bijugate shoot is therefore carrying a hazard of order one half per loss, with no route back.

Which makes three predictions, all cheaper to check than anything else in this thread.

Jugacy should be commoner at the base of a shoot than at the top, because the top has had fewer plastochrons in which to lose an organ but the base has had the whole season — and the loss is recorded above the point at which it happened, not below. A shoot that is bijugate early and spiral late is what an irreversible hazard applied node by node produces.

Jugate arrangements should be scarcer in the field than the rate at which they are initiated. If some fraction of apices set out bijugate, the fraction observed later is that fraction times the chance of having escaped a mid-front loss, which falls with age. Nothing here gives the initiation rate, so the prediction is comparative rather than absolute: the observed share should decline up a population’s age range.

And a shoot that has changed should carry the evidence. The wrecked stems settle at a whorl divergence the arrangement did not have — 23.44° or 156.56° at the coarsest rise, held constant — so the transition is not a gradual loss of symmetry but a step to a new angle. A specimen caught after one should show a sharp change in divergence at one node rather than a drift, and the node it happened at should be identifiable.

None of that needs a count. It needs somebody to walk up a stem noting how many organs sit at each node, which is the observation this essay has already argued is the cheapest in the thread — and the rate above is what makes it worth the walk rather than merely possible.

The caution that goes with it is the ordinary one. Three in five is the rate for this rule at this arrangement with a single organ removed from a swept set of offsets, and a real loss is neither uniformly placed nor always single. It is the right order of magnitude to plan an observation against and not a number to quote at a plant.

Why the underlying lattice does not surface

A reading, offered as one. The bijugate stem’s organs are placed against a neighbourhood, and the neighbourhood does not know it is two interleaved copies of anything. What it contains is ten short chains one way and six the other, and the rule’s profile is dominated by the organs at those two offsets. When the symmetry is broken the two copies stop being copies, and there is no longer any sense in which a smaller lattice is present to be fallen back onto.

Put the other way: the ordinary lattice underneath a bijugate one is a fact about how the pattern can be described, not about what any organ is placed against. A description does not have to survive an intervention.

What the experiment would look like on a plant

The translation is unusually direct here, which is the argument for doing it.

Find a shoot with two organs at every node — teasel and valerian are the textbook cases, and both are common. Count its parastichies; they will be even. Remove one primordium at a stated offset and watch what the shoot does next.

Three things are predicted, in decreasing order of how easy they are to see. The shoot should stop having two organs per node, permanently. The organs placed immediately after the cut should move if and only if the primordium removed was one of the most recent n, where n is the larger of the two counts. And removing either member of one node should give the same answer, which is a control the ordinary experiment cannot offer at all — two interventions that must agree, on the same shoot, decided by the pattern’s own symmetry.

The first of those needs no protractor and no counting after the fact. It is a yes-or-no question about whether a shoot still looks whorled, asked a few weeks later, and every account in which a whorled pattern is maintained by something other than the local arrangement predicts that it should recover.

What this does not say

It does not say the whorled rule is the rule plants use. It is the smallest extension of this collection’s own placement rule that can produce organs arriving in pairs, and what recommends it is that it produces the bijugate ladder the arithmetic predicts without being told to. Whether a meristem does anything like it is a different question.

It does not say bijugate stems are more fragile. The front is the larger count, as on an ordinary stem; the offsets that fail are in the middle of it, as on an ordinary stem. What is different is which property is lost, not how easily.

It does not say the repeat-unit account is finished. It says the account’s one sharp prediction — a block belonging to the underlying ordinary lattice — fails where it should have been strongest. An account that predicted the bijugate numbers for a different reason would be untouched by this, and nobody has written one.

And it does not say the symmetry could not return over a longer run. Three hundred organs after the cut it has not, at any offset, at any of the three rises. Longer runs are cheap and have not been done.

The check that would refuse it

Four assertions run when these figures are drawn.

The whorled rule has to produce a lattice with the rotational symmetry of its own jugacy, with both parastichy numbers multiples of it, and with a settled whorl divergence. If the extension quietly produced an ordinary lattice with a doubled count, everything after it would be about the wrong object.

The response has to be a run ending at the larger parastichy number when read modulo one k-th of a turn, with nothing past it — and the unfolded displacement has to reach past a quarter turn somewhere it has not moved. The second half is what makes the folding a correction rather than a convenience: if the raw numbers never misbehaved there would be nothing to fold.

Removing either organ of one whorl has to displace the next organ by the same amount, to within a degree. That is the symmetry showing up in the experiment, and a stem that failed it would not be bijugate whatever the counter said.

And every block reported has to belong to the bijugate pair and none of them to the ordinary lattice underneath. The check is made at the arrangement where the orbits are wide enough to be orbits, because at the two coarser rises there is no block to check — and reporting their periods as blocks would have put a two into the table at exactly the rung where the account under test predicts a two.

What links here

Computed from the collection, not written here: the essays that point at this one.

Shares its objects with

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

  • The pattern the cut leaves behind — both name ablation, attractor, counting blind, honest limits, jugacy, lattice, measurement, parastichy pair, the placement rule, whorl
  • A count with a factor in it — both name ablation, bijugate, counting blind, honest limits, jugacy, lattice, measurement, parastichy pair, rotational symmetry
  • A cut of two organs — both name ablation, honest limits, lattice, measurement, parastichy pair, the placement rule, refusal, self-correction
  • A wreck has a short list — both name ablation, attractor, counting blind, honest limits, lattice, measurement, parastichy pair, the placement rule
  • The stem that changed hands — both name ablation, attractor, counting blind, honest limits, lattice, measurement, parastichy pair, the placement rule
  • A front with no middle — both name ablation, honest limits, lattice, measurement, parastichy pair, the placement rule, self-correction

Named objects

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

AblationAttractorBijugateCounting blindChain countingHonest limitsJugacyLatticeMeasurementParastichy pairThe placement ruleRefusalRotational symmetrySelf-correctionWhorl