Packing and tiling
Why the average cell has six sides
Not because hexagons are efficient. Because Euler's formula leaves a tiling no choice — count the edges two ways and the mean comes out at six, whatever the cells would prefer. The efficiency argument is a different claim about a different thing.
Packing, measured four ways
The claim is that the golden angle packs best, and it is measurable. Read on the interior of a head, the two criteria about distance put the golden angle first among the angles near it and the two about cells are won by rational angles — which makes the claim half right, and makes the right half a statement about a class of angles. An earlier reading of the same four criteria, divided by the cells at the head's edge, said the opposite.
The gap that grows
A rational divergence angle develops an empty wedge that grows without bound as the head fills — a factor of 2.9 between two hundred primordia and sixteen hundred, and 3.8 from a hundred and fifty to two thousand. An irrational one does not. That is the division between rational and irrational angles that survives measurement.
Lewis's law wants disorder
Cell area rises linearly with side count — measured on cucumber epidermis in 1928 and quoted ever since as a property of packed tissue. It holds beautifully on a random point set, with a fitted constant of 1.64 against Lewis's 2. On a phyllotactic head it does not hold at all: the slope is 0.009, and area and side count are almost independent.
Two laws that want opposite tissue
Lewis's law and Aboav's relation are quoted side by side as properties of cellular tissue. Measured on the same two tilings they point opposite ways — the ordered head satisfies Aboav's with the textbook value of 1.18 and fails Lewis's completely; the random set does exactly the reverse.
The second moment is the measurement
The mean number of sides in a cellular tissue is six, and Euler's formula leaves it no choice — so it takes the same value on a golden-angle head, a whorled head and a set of random points. On heads of nine hundred organs the mean squared departure from six varies by a factor of eighty across the same three, and almost nobody reports it.
The six are the spirals
Label every contact between two cells in a seed head with the difference between the two nodes' placement indices. The labels are the parastichy numbers — 34, 55, 21, 89 — and the six sides Euler forces turn out to be about two from one family, one and a half from the next, and one each from two more.
The disorder is a staircase
Sweep the second moment of a head's side-count distribution across the divergence angle and it is not a curve. It is flat in stretches with sharp steps between them and narrow deep dips wherever a rational falls — so 0.253 is not the golden angle's number, it is the number of every angle from 137.47° to 137.54°.
A dip belongs to the head
At an exact rational the disorder halves when the head doubles, and the dip around it narrows by a factor of four. So which angles look ordered is set by how many organs were counted, and a head of three hundred cannot tell 138.4615° from 138.48° while a head of twelve hundred tells it from 138.4625°.
The background is not one sample
The dip in disorder at a rational angle is a comparison against a background, and the background was one measurement taken two tenths of a degree away. At 21/55 that lands seven thousandths of a degree from 13/34 — inside another rational's dip — and the comparison inverts. Fixed, the dip survives to a denominator of 89.
The width carries the denominator
The earlier work measured three denominators, found the dip's half-width falling as the square of the head size, and could not say whether its coefficient depended on the denominator. Six denominators say it does: the coefficient runs from 1,188 at q = 8 to 14,297 at q = 89, and dividing by q flattens a factor of twelve into a factor of two.
Four fractions with one denominator
The dip in a head's side-count disorder is as wide as 150·q/n², measured over six fractions — every one of them a Fibonacci convergent, which is the emptiest neighbourhood a denominator ever gets. So the law could be about the denominator or about how well the fraction approximates its neighbours. Four fractions of 55 at one head size settle it in one figure.
A width read off a staircase
Two fractions of the fourteen measured return a dip width that moves by a factor of two when the head size changes, where the others hold to three per cent. The cause is not their neighbourhood. It is that the disorder statistic changes only when the tessellation changes, so the curve a half-width is read off is a staircase, and a width narrower than the tread cannot be read at all.
A dip with no outer edge
The disorder of a head dips at every rational divergence, and how wide that dip is has carried a long argument. Reading the width as a level crossing has a resolution problem, and the obvious repair is to integrate instead. The integral reproduces beautifully across head sizes and never settles on a value, because there is nothing out there for it to settle against.
The window is the neighbour
An integral needs a limit, and this one has two conditions on it that pull opposite ways. It has to scale with the dip, so that two head sizes are comparable, and it has to stay clear of the next rational, which is a fixed distance in degrees. Between them there is no stretch where the answer holds still — and the limit that decides it is the crowding.
Fractions with the same neighbours
Every instrument this collection has for the width of a disorder dip has a free parameter set by how close the next rational sits — which makes a hypothesis about the neighbourhood untestable with any of them. The repair is not a better instrument. It is a set of fractions whose neighbourhoods are identical and whose denominators are not, and the arithmetic supplies twenty-four of them.
An interior that is nearly neutral
Give every cell a charge of six minus its number of sides and the total over a tessellated head is fixed by its own boundary, exactly, with nothing left over for the interior. On a golden head that freedom is spent on 264 exceptions among 1,631 cells which cancel to six.
The defects lie on rings
The cells in a seed head that are not hexagons are not scattered through it. Every one of 264 sits within a cell of a radius computed from the divergence angle alone, the radii are a factor of φ apart, and between two of them lie 422 consecutive cells without a single exception.
Every five is bound to a seven
A five-sided cell beside a seven-sided one is one object in a crystal and two exceptions in a tiling, and the phyllotaxis literature borrows the crystallographic word without measuring the binding. Measured against a seeded permutation null on a 2,400-organ head, every five in the interior shares a wall with a seven, and the loneliest one in the head is 0.927 cell spacings from the nearest.
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.
No cut-off makes them one
Two different relations on a head have both been called neighbour: the shortest index lags a count keeps, and a shared Voronoi wall. A cut-off that turns the first into the second exists for almost every cell taken alone, and for no whole head at any size.
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.
Two rankings, one list
An essay in this collection claimed that the four shortest index hops on a seed head and the four largest shares of its cell walls are the same four numbers in the same order, and called the correspondence exact. Measured again from the same points, the two lists hold the same four families and order them differently, and they order them differently in five of the six bands the head can be read in.
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.
Packing, measured against the interior
An earlier reading of these heads reported that no packing criterion singles out the golden angle and that three criteria give three winners. Every one of those readings was divided by a mean cell area that, on a head of 150 organs, was 38.8 where the interior's is π. Divided by the interior's own, the criteria about distance put the golden angle first of 72 angles and the criteria about cells go to rational ones.
One over root two
On the interior's scale a golden head's largest empty circle is 0.8435 of a spacing at every size from 150 organs to 2,000, because one triangle at its centre decides it. Everywhere else it is 1/√2 — a square cell at every ring where the lattice flips — and a closed form in the angle's continued fraction says that only noble angles hold it there.
The empty interval is the rings
No single cut-off on hop ratio turns the contacts a count keeps into the walls a tessellation draws, on any whole head at any size. Read cell by cell against the flip rings the divergence angle puts in closed form, every disputed cell lies within two thirds of a wall spacing of a ring, and with one spacing either side set aside a single cut-off between 1.430 and 1.444 serves every golden head from 900 organs to 9,000.
A second moment that goes to zero
The mean squared departure of a cell's side count from six separates a random tissue from a whorled head by a factor of eighty, on heads of 900 organs. Read at thirty-three head sizes it is exactly the share of cells on the defect rings of a spiral head and falls as one over the square root of the organ count, it falls as one over the count on a whorled head, and the factor is 25 at 300 organs and 677 at 8,000.