Phyllotaxis
the spirals are counted, not admired
Concept

Da Vinci's rule — where it appears

The claim that a branch's cross-sectional area is conserved at a fork, which is Murray's law with an exponent of two instead of three. Distinguishing the two on real junctions needs a range of daughter ratios that most measured trees do not supply.

Named by 20 essays across one field — each of them below, with the objects they name alongside it.

The exponent fitted from the junctions, rather than assumed. Sweeping k and asking where r₀ᵏ = Σ rᵢᵏ holds best gives 3.000 — Murray's 3, recovered rather than imposed.

Fitting the exponent

Assuming the exponent is three and reporting the error says how far the data is from that assumption. Fitting the exponent and reporting what it comes out as says what the network is doing — and an estimator has to be shown returning something other than three, or it is not a fit.

branching · Exponent The uninformative sample does not say so — it says something else. Above: how many junctions of a given asymmetry it takes to distinguish an exponent of 3 from an exponent of 2, at 2% measurement error on each radius. An even fork needs one; a junction whose small daughter is a twentieth of the large one needs 2195. Below: 50 junctions from each end of the range, on a synthetic tree built at exactly 3. The even forks return 3.01; the twigs return 1.69, with an interval no wider — because 47% of them measure as a parent thinner than its own larger daughter, and dropping those keeps only the half where the noise ran the right way.

Which junctions say anything

Da Vinci's rule and Murray's law differ by 12% at an even fork and by a tenth of a per cent at a twig. So one even fork settles which is right, and two thousand twigs do not — a factor of two thousand across a tree, decided entirely by the shape of the junction and not by how carefully it is measured.

branching · Exponent The uninformative sample does not say so — it says something else. Above: how many junctions of a given asymmetry it takes to distinguish an exponent of 3 from an exponent of 2, at 2% measurement error on each radius. An even fork needs one; a junction whose small daughter is a twentieth of the large one needs 2195. Below: 50 junctions from each end of the range, on a synthetic tree built at exactly 3. The even forks return 3.01; the twigs return 1.69, with an interval no wider — because 47% of them measure as a parent thinner than its own larger daughter, and dropping those keeps only the half where the noise ran the right way.

A sample that is confidently wrong

Fifty lopsided junctions from a tree built at an exponent of exactly 3 return 1.7, with an interval that excludes 3 and excludes 2 as well. The sample carrying almost no information does not give a wide answer — it gives a narrow wrong one, and the cause is a selection nobody applies on purpose.

branching · Exponent A tree built at an exponent of 3, measured band by band. Six bands of daughter ratio, 300 junctions in each, all from trees built at an exponent of exactly 3 with the same 2% measurement error. The median implied exponent falls from 3.00 at an even fork to 1.58 at a twig, and the share of junctions that have any exponent at all falls from 100% to 53% over the same range. The rule across the top is a single fit over 200 junctions spanning the whole range: 2.85. A mixed sample is safe because least squares already weights by leverage — 42% of it sits in the most symmetric band and 0.06% in the twigs. The sample that is not safe is the one a person can reach.

The band decides the answer

A fit over a whole tree's junctions returns the exponent the tree was built at, even though most of its junctions are from bands that on their own return 1.6. Least squares is already weighting by leverage. The dangerous sample is not the mixed one — it is the one a person can reach.

branching · Exponent A tree built at 3, measured to 2%, reads 2.957 on the informative band. The exponent recovered from 100 junctions of a tree built at exactly 3, against the relative error placed independently on the parent and on both daughters — half a per cent is a machined section under a microscope, one to two per cent is callipers on a clean branch, five is a branch that is not round, ten is a radius read off a photograph. Each line is a band of daughter ratio and the whiskers are the central 90% of 300 replicate samples. Every band is displaced downward at every error and never upward: at 2% the informative band read 2.957 and the informative band read 2.957. Below, the same rows with the displacement and the spread drawn as separate bars, because only one of them falls when more junctions are measured.

The exponent an error moves

Every real measurement of a branch radius carries error and no synthetic tree does, so the question is what a symmetric error does to a fitted exponent. It does two things — a bias and a spread — and the bias runs downward at every error level and in every band, by an amount derivable from the daughter ratios alone.

branching · Exponent error One junction's bias moves 1.36-fold across the range its leverage moves 308,352-fold, at an exponent of 3. The bias a single junction of daughter ratio γ contributes to a least-squares fit, as a multiple of the squared measurement error, drawn against the leverage that junction carries — both computed from the expansion about a true exponent of 3 rather than fitted to anything. Across the whole range from an even fork to a twentieth the bias moves by a factor of 1.36 and the leverage by a factor of 308,352, and the uninformative junction contributes the larger share: 6.00σ² at γ = 0.05 against 4.50σ² at an even fork.

The fragile junctions are the informative ones

That is the obvious worry once the radii are uncertain, and it is false. Across the whole range of asymmetry a junction's contribution to the bias moves by a factor of 1.36 while its leverage moves by a factor of 308,352, so the junction that says nothing damages the answer as badly as the one that says everything — and a sample is spoiled by counting rather than by weight.

branching · Exponent error Between 3% and 5% of radius error, no sample size answers — 50 junctions among them. One row per error level. The pale bar is the sample sizes whose interval is narrow enough to state a claim from — half-width under ±0.25 and excluding 2 — and it starts where precision arrives. The second bar is the sample sizes whose interval still contains the 3 the tree was built at, and it ends where the displacement overtakes the width. Where the two overlap there is a usable window; at 5%, 7%, 10% they do not overlap at all, so below 50 junctions the answer is too wide to state and above 30 it no longer contains the truth.

The window that closes

The spread of a fitted branching exponent falls as the reciprocal root of the sample and its displacement does not fall at all, so there is a count past which every further junction buys confidence and no accuracy. Between three and five per cent of radius error the count arrives before the answer does, and no sample size both states a claim and contains the truth.

branching · Exponent error Murray's law and Da Vinci's become one measurement at 12% of radius error on the informative band. Two trees measured the same way: 50 junctions of daughter ratio 0.6–1, 300 replicate samples at each of 15 error levels, one tree built at exactly 3 and one at exactly 2. Each shaded band is the central 90% of the recovered exponents. Both run downward, but the tree at 3 runs down faster — -113σ² against -29σ² — so the two close on each other. At 11% they are still apart; at 12% the bands overlap and one study's answer could have come from either tree; at 20.5% the means cross, and above it a tree built at 3 measures lower than a tree built at 2.

Where three and two become one

Two trees, one built to obey Murray's law and one to obey Da Vinci's, are measured through the same fifty junctions with the same instrument. At twelve per cent of error on each radius the two answers overlap, and above twenty and a half the tree built at three measures lower than the tree built at two.

branching · Exponent error The three weights at an exponent of 2 and 3: one closes a triangle and one is a straight line. A branch point minimising a weighted sum of three lengths has an interior solution only when the three weights close a triangle, and the weight on a segment here is its own cross-section. At an exponent of 3 the three areas clear that condition by 0.4126, and the triangle they close is what the two fork angles are read off. At an exponent of 2 the parent's area is exactly the daughters' areas summed — that is what area conservation says — so the slack is -4.44e-16, the triangle collapses onto a line, and the fork closes to 0.0000 degrees. Leonardo's rule does not predict a different angle here; it predicts no angle.

A rule that predicts everything

Leonardo's rule says a fork conserves cross-section, and cross-section is exactly the weight the branch point is minimised against. So the three weights land on the boundary of the triangle inequality, the cosine comes out at one to the last bit, and the rule predicts no angle at all — and the free constant its own derivation leaves behind then walks the prediction across every angle a fork could have.

branching · Fork angle What a swelling does to the exponent read from the informative band, by where the swelling is. 50 junctions of daughter ratio 0.6–1, built at exactly 3 and read with no random error, but with one or more radii measured fat. A parent read fat lowers the reading: 1% gives 2.864, 3% gives 2.631, 10% gives 2.075, and at 11.3% the tree reads Da Vinci's 2. Daughters read fat raise it: 1% gives 3.150, 3% gives 3.499; from 8% some junctions have a daughter measured wider than their parent, which no exponent balances, and the line stops. All three radii read fat by one factor return 3.000 at every swelling — the unswollen reading exactly.

A swelling at the fork

A branch thickens where it forks, so a parent measured just below a junction and daughters measured just above it carry three different amounts of the same swelling. A swelling that fattens all three alike moves a fitted exponent by exactly nothing. A parent read one per cent fat moves it by as much as 3.6 per cent of random error on every radius, in a sign known in advance, and a tenth of a radius turns a tree built at Murray's three into one that reads Da Vinci's two with no noise at all. Added to the noise, it does not bring the two rules together any sooner: the two errors do not add.

branching · Exponent error Adding error to carry a fitted exponent back to none, at 12%. At 12% of error on every radius each replicate is refitted with more error added at four levels, and the dots are the means over 300 replicates, the tree at 3 above and the tree at 2 below. Uncorrected they read 2.035 and 1.676. Carried back to no error through the mean points, the tree at 3 reads 2.695 by a quadratic curve, 2.380 by a linear curve, 2.914 by a rational curve; the tree at 2 reads 1.961 by a quadratic one, 1.877 by a linear one, 1.982 by a rational one. The curve carried back is a choice the method does not make.

A correction that keeps the overlap

The duel between a tree built at Murray's exponent and one built at Da Vinci's ended by saying the displacement is the geometry, and that no better estimator removes it. Correcting every replicate by simulation-extrapolation removes 92 per cent of the tree at three's displacement at five per cent of error and 68 per cent at twelve, and the error at which the two means cross leaves the measured range altogether. It pays in spread — the corrected readings are twice as wide at twelve per cent — so the error at which the two trees' intervals overlap does not move. Of the duel's two numbers, the inversion was the estimator's and the overlap is the question's.

branching · Exponent error Measured radii against the tips each branch carries, at 12% of error on every radius. One replicate of a 50-junction tree, 101 segments, every radius measured with 12% of relative error and drawn against the number of tips that segment carries. The count has no error in it, so the slope is not displaced: the tree built at 3 gives a slope of 0.3404, an exponent of 2.937, and the tree built at 2 a slope of 0.5071, an exponent of 1.972. On the same measured radii the junction-by-junction fit reads 2.218 and 1.736.

A count carries no error

Fitting r₀ᵏ = Σrᵢᵏ junction by junction puts a measured radius on both sides of every equation, and at twelve per cent of error a tree built at Murray's three and one built at Da Vinci's two stop being told apart, however the fit is corrected. Fit the same measured radii against the number of tips each branch carries instead — a count, which nobody measures with error — and the two trees read 2.996 and 1.997 at twelve per cent and 3.015 and 2.001 at thirty, never overlapping. The twelve per cent belonged to the junction fit, not to the tree. The count fails in its own way, and the way is stated.

branching · Exponent error One tree sized for flow and for stress, with each branch 2^(−1/2) the length of its parent. The same symmetric tree, 8 generations deep, each generation's branches 2^(−1/2) the length of the one before and turned 30° at every fork, sized two ways and drawn to one trunk width. On the left each branch's radius cubed is proportional to the tips it feeds — Murray's flow rule — and every junction conserves r³. On the right each branch is sized so that the same load on every tip bends it to the same stress at its base, radius cubed proportional to the sum of its lever arms to its tips; its trunk junction conserves r to the power 1.967, its outermost junctions 1.349, against a deep-tree limit of 2.000. The two trees thin at different rates from the same trunk.

A cube law with a lever arm

Murray's exponent of three comes from moving fluid for the least work, and Da Vinci's two has had no derivation here, only the name of the mechanical answer. Size every branch so that the same wind on every tip bends it to the same stress, and a junction conserves r to the power 3/(1 + log₂(1/λ)), where λ is how much shorter each branch is than its parent. A crown that fills a plane gives exactly two; halving lengths gives one and a half; no shortening gives three. Murray's flow rule gives three at every λ, so the lengths of a tree's branches say which mechanism sized it.

branching · Murray The fork angle Da Vinci's rule predicts, against the size of the fork, in one tree. A tree has one value of the constant Da Vinci's rule leaves free, so a fork's share of it falls as the square of the fork's size and the angle the rule predicts changes with size. For even forks it is 119.1° at a relative size of 0.1, 101.7° at a relative size of 0.5, 74.9° at a relative size of 1, 44.6° at a relative size of 2, 9.6° at a relative size of 10, where Murray's rule gives 74.93° at every size; for daughter ratio 0.5 it is 118.9° at a relative size of 0.1, 101.3° at a relative size of 0.5, 77.6° at a relative size of 1, 48.9° at a relative size of 2, 10.9° at a relative size of 10, where Murray's rule gives 77.58° at every size. From a total of 100° to 20° at an even fork is a factor of 8.93 in radius.

One constant for every fork

Da Vinci's rule leaves a free constant in the cost that sets a fork's angle, and running it over its range walks the predicted angle from nothing to 120 degrees, through Murray's 74.93. So no single fork can refute the rule. But the constant is one number for a whole tree, and a fork's share of it falls as the square of the fork's size — the constant is a radius axis. A tree spanning a factor of ten in radius must show forks from 29.4 degrees at its biggest to 111.6 at its smallest, a spread wider than one fork's flatness can hide, while Murray's angle is the same at every size.

branching · Fork angle The exponent read from a tree that has lost tips, tips lost one at a time. The 50-junction tree built at 3 and at 2, 12% of error on every radius, 300 replicates of loss and measurement at each level. Shaded: the central 90% of the exponent read against the tips still there. Dotted: the mean read against every tip ever grown. Solid: the junction fit's mean over the junctions that survive. The count's two intervals are still apart with 70% of the tips gone, reading 2.366 and 1.576, and overlap by 80%. The junction fit's intervals overlap by 20%. At the heaviest loss drawn, 90%, the scars still read 2.998 and 1.997.

A count that has lost tips

Radii read against the tips each branch carries keep Murray's three apart from Da Vinci's two where junction fits cannot, because a count has no measurement error in it. A count of the tips a tree has is not a count of the tips it grew. Losing them lowers both trees' readings by one factor that belongs to the losses and not to the rule, so the count stops being right long before it stops telling the trees apart: on fifty junctions at twelve per cent of error, to seventy per cent of the tips lost one at a time, and only to about a quarter lost in whole limbs. Counting scars repairs single losses exactly. Nothing countable repairs a shed limb.

branching · Exponent error The fork angle the transport cost predicts on a tree sized by stress, against the size of the fork. A tree sized so that equal loads on its tips bend every branch to one stress conserves rᵖ with p set by how much shorter each branch is than its parent, λ. On such a tree the cost that fixes a fork's angle carries one constant, entering a fork of size s as s^(2p − 6), so the predicted angle changes with size unless p is three. With the constant set so that a fork of relative size one opens at Murray's 74.9°: At λ = 0.707 (p = 2.000) an even fork opens at 119.1°, 111.6°, 74.9°, 29.4°, 9.6° at relative sizes of 0.1, 0.32, 1, 3.2 and 10; at λ = 0.74 (p = 2.091) an even fork opens at 114.5°, 106.4°, 74.9°, 41.1°, 30.1° at relative sizes of 0.1, 0.32, 1, 3.2 and 10; at λ = 0.794 (p = 2.250) an even fork opens at 106.3°, 97.6°, 74.9°, 53.9°, 46.4° at relative sizes of 0.1, 0.32, 1, 3.2 and 10; at λ = 0.87 (p = 2.498) an even fork opens at 92.7°, 85.5°, 74.9°, 66.0°, 61.5° at relative sizes of 0.1, 0.32, 1, 3.2 and 10; at λ = 0.95 (p = 2.793) an even fork opens at 78.4°, 76.6°, 74.9°, 73.4°, 72.2° at relative sizes of 0.1, 0.32, 1, 3.2 and 10. Murray's rule opens every fork at 74.9°.

Forks on a tree sized by stress

Da Vinci's rule predicts that a tree's forks open wider as they get smaller, because the constant in its angle cost is one number for a tree and enters each fork scaled by its size. A tree sized for equal bending stress conserves an exponent set by how much shorter each branch is than its parent, and on such a tree the same constant enters each fork as its radius to the power 2p − 6. The trend survives at every length ratio short of one and shrinks with it: 105 degrees a decade of radius for a crown filling a plane, 83 at a length ratio of 0.74, 53 for a crown filling a volume, 13 at 0.9. Only a crown shortening about as fast as a volume-filling one fans wider across a tenfold range than one fork's flatness can hide.

branching · Fork angle One planar crown sized for a load on its tips and for the weight of its own wood. The same symmetric crown, 9 generations deep, each branch 2^(−1/2) the length of its parent and turned 30° at every fork, sized so that every branch is bent to one stress, drawn to one trunk width. On the left the load is on the tips and the trunk junction conserves r to the power 1.980; on the right the load is the weight of the wood, found by iterating the radii until they stop moving, and the trunk junction conserves r to the power 0.969. A crown sized for its own weight thins much faster from the trunk, because a branch's weight grows with the square of its radius.

A crown that carries its own wood

Sizing every branch so that equal loads on the tips bend it to one stress gives a crown filling a plane Da Vinci's exponent of two. Move the load onto the wood and the sizing becomes a fixed point, because a branch's load now depends on the radii being solved for. Under the wind on its wood a planar crown still conserves two, but only as a limit its trunk is two tenths short of at fifteen generations. Under its own weight it conserves one — radius rather than area, the stress-similarity law that radius goes as length squared — and a crown carrying leaves and wood reads the leaves' two near its twigs and the wood's one at its trunk, with the handover set by how much of the trunk's load the wood carries.

branching · Murray Sizing for equal bending and sizing for equal stress cross at one length ratio. Equal stress holds r³ against the sum of a load's arms and gives 3/(1 + ℓ); equal deflection holds r⁴ against the sum of the arms squared and gives 4/(1 + 2ℓ), with ℓ = log₂(1/λ). Setting them equal gives 3(1 + 2ℓ) = 4(1 + ℓ), whose only root is ℓ = 1/2 — the crown that fills a plane, λ = 0.707107 — and there both are exactly two. Below that ratio the stiffness rule reads the lower exponent of the two and above it the higher, so the two criteria size the same crown at one length ratio in the whole family and it is the one Da Vinci's rule names.

A crown sized for how far it bends

Stress is one criterion for sizing a branch and stiffness is another. Holding every branch to the same deflection as a share of its own length sizes r to the fourth against the sum of each load's arm squared, where equal stress sized r cubed against the arm, and the junctions of a deep crown then conserve 4/(1 + 2·log2(1/λ)). A single cantilever under its own weight comes out at radius as length to the three halves — McMahon's elastic similarity, fitted here rather than assumed — against the square that equal stress asks for. And the two criteria agree at exactly one length ratio out of the whole family: λ = 2 to the minus a half, the crown that fills a plane, where both give exactly two.

branching · Murray Three ways of sizing a crown, and the exponent each one conserves. Murray's flow rule sizes r³ against the tips a branch feeds and conserves three at every length ratio, reading nothing of the lengths at all. Equal bending stress conserves 3/(1 + ℓ) and equal deflection 4/(1 + 2ℓ), where ℓ = log₂(1/λ). So an exponent measured on a tree names a rule only with a length ratio beside it, and even then not everywhere: the stress and stiffness curves meet at λ = 0.7071, the stiffness curve passes three at λ = 0.8909, and the stress curve reaches three only as the branches stop shortening.

Three rules, one exponent

A measured branching exponent is quoted as evidence for a sizing rule, and it cannot be. Murray's flow rule conserves three at every length ratio and reads nothing of the lengths at all; equal stress conserves 3/(1 + l) and equal deflection 4/(1 + 2l), where l is log2(1/lambda). So an exponent names a rule only with a length ratio beside it, and even then not everywhere: of ninety-six length ratios between 0.3 and 0.99, thirteen have two rules within five hundredths of each other at a precision of 0.05, in three bands with three different reasons — stress against stiffness where they cross at the planar crown, stiffness against flow where the stiffness curve passes three at 0.8909, and stress against flow only as the branches stop shortening.

branching · Murray Fork angles on crowns sized by the larger of flow and stress, handing over at six different generations. Thirteen-generation crowns at a length ratio of 0.707, every branch sized by the larger of flow and stress, twigs at the radius the transport cost prefers, with the two radii equal at generations 3 to 8. Handing over at 3, the forks from the trunk open at 38, 51, 67, 75°; handing over at 4, the forks from the trunk open at 27, 38, 51, 67, 75°; handing over at 5, the forks from the trunk open at 19, 27, 37, 51, 67, 75°; handing over at 6, the forks from the trunk open at 13, 18, 26, 36, 50, 66, 75°; handing over at 7, the forks from the trunk open at 8, 12, 17, 24, 35, 49, 66, 75°; handing over at 8, the forks from the trunk open at 4, 6, 10, 15, 22, 32, 46, 65, 75°. Every fork from the handover outward opens at Murray's 74.9°. No fork on any of the six opens wider than that, and none turns back: the angle rises through the stress-sized generations and stops.

A trend that stops at Murray's angle

A crown sized by whichever of flow and bending stress asks for the thicker branch is sized by stress at its trunk end and by flow at its twigs, and the twigs fix the constant that was free in the fork-angle prediction: a twig at the radius the transport cost prefers spends exactly half its upkeep on pumping. On such a crown the fork angle does not change sign at the handover. It rises through every stress-sized generation, meets Murray's 74.93° at the handover and stays there, and no fork anywhere opens wider. The trend turns back only when the twigs are thinner than the cost wants — past a pumping share of (λ^(−2/3) − 1)/(1 − λ^(4/3)), 0.70 at the planar crown and closing on one half as branches stop shortening.

branching · Fork angle

Named alongside it

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

Branching exponentMurray's lawHonest limitsMeasurement errorBiasClaim testingDiscriminationPredictionClosed formEvidenceFittingIdentifiability

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