Phyllotaxis
the spirals are counted, not admired
Concept

Morphospace — where it appears

A parameterisation of possible forms in which a real organism is a point and the unoccupied regions are the question. Raup's three numbers put nearly every coiled shell that has existed into one, and reading its empty regions as forbidden rather than merely unvisited is where most over-reading of it happens.

Named by 7 essays across one field — each of them below, with the objects they name alongside it.

A shell section at W = 2.40, D = 0.42. W·D = 1.01, so the whorls are free of each other — an evolute shell, like a ram's horn or a planispiral ammonite. Both are things animals grow.

Raup's three numbers

Nearly every coiled shell that has ever existed is a point in a three-dimensional space — how fast the whorl expands, how far the opening sits from the axis, how far it travels along it. That is a remarkable compression, and the most useful line in the space is the one where the whorls come apart.

shells · Raup The boundary located at 481 expansions, against D = 1/W. 481 expansions log-spaced across the range, each boundary found by bisecting the two drawn circles to two hundred steps rather than read off a grid. The hyperbola D = 1/W lies on the located curve to 2.220e-16 over the whole of it — the last bit of a double — so the textbook line is the boundary and not an approximation of it. The closed form and the bisection agree to 2.81e-15 at worst over all 3,200 located boundaries, which is what makes a residual of zero a reading.

The line was already exact

The boundary between shells whose whorls run into one another and shells whose whorls run free is quoted everywhere as D = 1/W, and a survey designed to measure how far off it sits found that it is not off at all. Located by bisecting the drawn circles at 481 expansions, the residual is 2.2 × 10⁻¹⁶ — the last bit a double holds, over the whole range.

shells · Morphospace The site's hero shell comes free at a translation of 1.1066, and a slower one at 10.49. Setting the located boundary to zero and solving leaves T_free = √W/(W−1), drawn here across the whole expansion range on logarithmic axes. Above the curve the shell is free at every distance from its axis and the contact region has left the plane rather than merely shrunk in it; a slowly expanding shell at an expansion of 1.1 needs 10.4881 turns of translation to buy that and one expanding twentyfold needs 0.2354. The tower is what buys a shell the right to coil close to its own axis, and the faster it expands the less tower it takes.

What a spire buys

Translation along the coiling axis enters the contact boundary as its square, so a shell that has only just begun to walk along its axis has not moved the boundary at all. It is also strictly one-way, and it has a threshold above which no distance from the axis whatever puts the whorls in touch.

shells · Morphospace Two measures of one boundary at W = 2.5: overlap slope 1, buried area slope 1.4999. The linear overlap and the buried area, against how far below the boundary the shell sits, both axes logarithmic. The buried fraction rises as 1.499945 — three halves, the lens between two nearly tangent circles — so at a ten-thousandth below the line 0.0011410 per cent of the whorl is under its successor and at 0.30 below it 86.83 per cent is. The linear overlap over the same range is exactly 2.5 times the distance below and falls straight to zero, so the two measures disagree about whether the boundary is sharp and the area is the one that answers the question. A shell just inside the boundary is not a different kind of shell; one well inside is.

A boundary with no edge

Two continuous measures cross the line where a coiled shell's whorls begin to touch, and they disagree about whether it is sharp. One falls to zero as a straight line and makes the line a kink; the other leaves it as a three-halves power and makes it a tangency.

shells · Morphospace One geometry, six boxes: 4.642 per cent to 52.81 per cent forbidden. The share of each box that the coiling geometry excludes, across six boxes that differ only in where their edges were put and in how one axis is sampled. The spread is a factor of 11.4, from 4.642 per cent in the widest box to 52.81 per cent in the tightest, and sampling the same two decades of expansion geometrically rather than uniformly multiplies the answer by 4.59 on its own. The forbidden area is the integral of 1/W and grows as a logarithm while a box grows as a line, so the fraction has no value of its own to quote.

A fraction of nothing

Six boxes differing only in where their edges were drawn give between 4.64 and 52.81 per cent for the same geometry, and sampling one axis geometrically rather than uniformly multiplies the answer by 4.60. The share tends to zero as the box widens, because the region under a hyperbola is a logarithm and a box is a line.

shells · Morphospace An axial section of a spire at W = 2.4, D = 0.3, T = 1, with the angle that decides contact. The discs where the plane holding the axis cuts three whorls, on both sides of the axis. Every disc subtends the same half-angle from the apex, γ = 17.065°, about the line of the disc centres at β = 33.024° from the axis, so the envelope's apical angle is 100.178° whatever the expansion. Whorls with that angle touch below W = 1.8307, and at 2.4 they run free. Successive discs on one side are 2.4 times farther from the apex, and one disc gives back D = 0.3000 and T = 1.0000.

One angle decides contact

Seen from the apex of its coiling axis, every whorl of Raup's shell subtends the same half-angle, and two whorls touch exactly when the sine of that angle exceeds (W − 1)/(W + 1) — with no disagreement against the drawn discs at 400,000 random shells. Two of the three numbers enter only through the angle and the third only through the threshold, which decides which picture of a shell can answer the question: a spire's outline carries no W, a plan carries no T, and an axial section carries all three.

shells · Morphospace An ellipse twice as tall as it is wide at W = 2.4, T = 0.5, beside a circle at T = 0.25. Each panel is an opening, in colour, and the same opening one whorl on, 2.4 times larger about the apex, drawn at the axis distance where the two just meet. An ellipse twice as tall as it is wide at a translation of 0.5 meets at D = 0.391257; a circle at a translation of 0.25 meets at D = 0.391257, the same number, because stretching the axis by 1/2 turns the ellipse into the circle and 0.5 into 0.25.

The fourth number divides the third

Every boundary on Raup's cube was located for a circular opening, and three essays ended on the same hedge: the numbers would move with a differently shaped aperture by an amount nothing had measured. Measured on the drawn outlines of eleven openings, the boundary with no translation does not move at all for any convex opening symmetric about the plane of coiling; an ellipse's height divides the translation and does nothing else; the square law in the translation belongs to a round tip; and a turned opening frees ground only in the D a plan reads.

shells · Morphospace

Named alongside it

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

Honest limitsModel scopeWhorlEvoluteInvoluteGrowth factorParameter spaceAxis translationClaim testingClosed formResolutionSpire

All concepts