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The Fundamental Theorem of Phyllotaxis revisited

The Fundamental Theorem of Phyllotaxis revisited ( July 2024): This paper has now been superseded and, whilst still mathemati- cally correct, should be considered obsolete. Jean’s ‘Fundamental Theorem of Phyllotaxis’ (Phyllotaxis: a systematic study in Plant Morphogenesis, CUP 1994) describes the relationship between the count numbers of observed spirals in cylindrical lattices and the horizontal angle be- tween vertically successive spots in the lattice. It is indeed fundamental to ob- servational studies of phyllotactic counts, and especially to the evaluation of hy- potheses about the origin of Fibonacci structure within lattices. Unfortunately the textbook version of the theorem is incomplete in that it is incorrect for an important special case. This paper provides a complete statement and proof of the Theorem. Note This paper was first uploaded to arXiv in 2012, and the counterexample here to Jean’s theorem as he stated it remains valid. Since that time Infang Publishing has pub- lished my book-length treatment of Mathematical Phyllotaxis, ISBN 0993178960. This book puts the (minor) signican fi ce of the problem in more context and su- persedes this paper, which should be considered obsolete – Jonathan Swinton, July 1 Introduction Mathematical phyllotaxis is the study of the patterns that appear in two-dimensional cylindrical lattices, given particular motivation by the striking appearance of high Fibonacci numbers in a range of biological settings such as the spirals on a sun- flower [6]. Although static analyses of lattices cannot in themselves explain the ap- pearance of these numbers [12], they are essential both in relating what can actually arXiv:1201.1641v3 [q-bio.TO] 9 Jul 2024 be biologically observed to hypothesises about the underlying order, and in forming a basis for dynamical models of lattice formation [8, 4, 10, 2] that can, it is claimed, explain Fibonacci numbers and related structure in biological form. More specifi- cally, a phyllotactic theory of lattices creates a model for which lines in the lattice are most likely to be remarked on by a human observer. In the case of the sunflower or the fir cone, these lines may be those which join adjacent points in the lattice, which may be defined in different ways as contact parastichies [9, 6] or principal paras- tichies [12]. A slightly more general idea is to identify those pairs of lines that wind in opposite directions as opposed parastichies, or alternatively those lines which can be thought of as characterising the lattice, which were defined as generating parastichies by Turing [12] or equvalently as visible parastichies by Jean [6]. Jean presents the most complete description in the literature of mathematical phyllotaxis in his textbook [6], and deserves considerable credit both for innova- tion and a substantial work of integration, bringing together a range of biological datasets and historical mathematical approaches. One major contribution is what he calls the Fundamental Theorem of Phyllotaxis which he attributes in a special case to Adler [1]. The basic idea of the theorem is a very useful conceptual one. Cylindrical lattices can be, for these purposes, completely characterised by the an- gle of rotation between successive points called the divergence. If a lattice is seen to posses a specic fi pair of generating and opposed parastichies, characterised in a natural way by a pair of integers, then there is a constraint on the allowable values of the divergence. The theorem shows that for any pair of integers not both equal to 1 there are exactly two intervals of nonzero width on which the divergence will create the required generating and opposed parastichies. Unfortunately, however, in the form stated by Jean the Theorem needs modification in a range of special but important cases. The goal of this note is to restate the Theorem completely. First we give the necessary background about two-dimensional cylindrical lattices, and give a characterisation of which divergence values correspond to a given generating pair. After restating the Fundamental Theorem in Jean’s formulation and demonstrating a counter example, we then reprove a suitably corrected Theorem. Unbeknown to Jean and Adler, Turing had also considered very similar problems, but this work was unpublished at his death in 1954 [11]. It remained accessible but obscure in the Turing Archive in King’s College Cambridge, until being published in his collected works in 1992 [12], well after the relevant papers of Jean and Adler. As a secondary aim, this paper points out the ways in which Turing anticipated the later, more widely known work. 2 Background This section contains a number of statements without proof that are fairly obvi- ous on examining a diagram. They can be made rigorous by eg the use of congru- ences [12]. We consider a cylinder of circumference 1 and extending infinitely in the vertical direction, with an origin and coordinates (x , z ), 0 ≤ x ≤ 1. For any 0 ≤ d ≤ 1, we can construct a lattice d by rotating by an angle 2πd around the cylinder from the origin 2 and rising by z = 1, and repeating. This creates the set of points (x , z ) = (d , m) where m is any integer and d = md − [md ] and [x ] is the nearest integer to x , so 1 1 1 1 that − ≤ d ≤ . Since taking [ ] = 0 and [ ] = 1 map to the same point on the 2 2 2 2 cylinder we will allow the function [x ] to take the multiple values 0, 1 at the point x = . We call (d , m) the point ℓ , and from now on assume m ̸= 0. The vertical m m component is called the rise. By construction we have excluded lattices with more than one point at each rise. More generally if there are J such points spaced equally around the cylinder we would describe the lattice as having Jugacy J , but we restrict to J = 1 here. A parastichy of order m is the infinite line through the origin and ℓ with slope d /m. There are two possible lines on the cylinder through 0 and ℓ corresponding m m to winding in opposite directions and this choice of slope is equivalent to choosing the line that traverses the smallest x distance between 0 and ℓ , leaving an ambi- guity when [md ] = . The portion of this line between 0 and m defines the vector m, again with an an ambiguity when [md ] = . An m-parastichy is a member of the family of m distinct lines containing the origin-parastichy of order m and the paral- lel lines to it through the points 1, . . . , m − 1. If a point ℓ is on an m-parastichy then so is ℓ . See Figure 1. p+m A pair of (parastichy) numbers (m, n) define a pair of points ℓ , ℓ and vectors m n m,n. (m, n) is opposed if d /m and d /n have opposite sign. In the case when m n [md ] = we define the parastichy pair ( m, m) as the pair combining each of the choices of direction around the cylinder. There remains an ambiguity when [md ] = 1 1 or [nd ] = and m ̸= n which could be resolved by a specific choice of direction 2 2 although it is not of significance subsequently. There is a natural relationship between a cylindrical lattice and a corresponding periodic lattice in the plane, and the m-parastichies also dene fi an infinite family of m-parastichies in the plane lattice. A pair is generating for d if it generates the lattice in the plane in the sense that every point can be expressed as a vector sum vm+ un for integer u, v . It is necessary, but not sufficient, for a generating pair (m, n) to be coprime for if they have a common factor k all rises, including 1 must be a multiple of k . This is effectively the definition given by Turing [12], and identical to the visible pair dened fi by Jean [5, 6] in a number of different equivalent ways. Since Jean [6] gives no proof of that equivalence we give it in the Theorem below which also establishes the identity with the Turing definition, and in the process modify some of Jean’s denitions fi for extra precision. I have chosed to stick with Turing’s word generating over Jean’s visible for these identical concepts as I think the latter word carries confusing connotations in the identification of parastichy counts. We make use of the determinant ∆ of a pair (m, n), defined as mn ∆ (d ) = [nd ] m − [md ] n mn = (nd − d )m − (md − d )n n m = nd − md , m n Note that the denition fi is such because any non collinear vectors would generate the lattice in the cylinder. 3 54 Figure 1: (4,5) is a generating and opposed pair for the cylindrical d = 17/72 lattice. The parallelogram defined by the pair tiles the lattice and every lattice point is at a vertex of one of the parallelograms; the edges of the parallelograms form the paras- tichy lines. The blue lines highlight the family of 4-parastichies and the red lines the family of 5-parastichies. 4 1 1 except for the special case ∆ ( ) = m (where we have picked [ ] = 1 in the [nd ] mm 2 2 and [ ] = 0 in the [md ]). Theorem 1. (Compare Theorem 4.2 of Jean [6].) The following are equivalent 1. The pair (m, n) is generating in the lattice d . 2. The pair (m, n) has a point of the lattice d at every intersection of the lines of the pair. 3. The points 0, ℓ , and ℓ form a nondegenerate triangle which contains no m n other point of the lattice d internally. 4. (m, n) satisfy |∆ (d )| = 1. mn Proof. If a pair of vectors are collinear in the plane, they cannot be generating. If they are not collinear, the parallelogram formed by any pair can be used to tile the cylinder. This tiling will contain lattice points exactly at the vertices of each parallel- ogram iff the pair is generating, because if it has an internal point it must be a nonin- teger sum of the pair. This shows 1⇔3. Moreover the tiling produces the parastichy families of order m and n, so these must always intersect at a lattice point iff the pair is generating. This shows 1⇔2. A pair is generating iff it can express the unit vector (0 ≤ d < 1, 1) as a sum of m and n in the plane. In plane coordinates, we have [nd ]m− [md ]n = ([nd ] d − [md ] d , [nd ] m − [md ] n) m n = ([nd ] (md − [md ])− [md ] (nd − [nd ]) ,∆ ) mn = ∆ (d , 1) mn If ∆ = 1 we are done, and if ∆ = −1 we take the combination [md ]n− [nd ]m of mn mn opposite sign, so if |∆ | = 1 then (m, n) is generating. This shows 4 =⇒ 1. mn To prove 1 =⇒ 4, the central idea (of Jean and Adler and Turing) is to continue the m and n parastichies away from the origin until they cross again, so first we have to dispose of the case when the two parastichies are parallel. If m and n are paral- lel, then they are not generating, and moreover d /m = d /n so ∆ = 0 and con- m n mn versely. If they are not parallel then in the plane the parastichy of order m through (0, 0) and (d , m) and and the parastichy of order n through (1, 0) and (d , n) must m n meet at the point µ ¶ md nd mn n m = − 1, . ∆ ∆ ∆ mn mn mn If (m, n) is generating this must be a point of the lattice and so have rise equal to ′ ′ ′ both k m and k n for integer k , k , so n = k∆ and m = k ∆ . But since m and n mn mn are coprime, |∆ | = 1. mn There are close connections with the theory of Farey sequences, as mentioned in Jean [6] and in more detail in Jean [5], which can be exploited to give different versions of this proof, but we do not pursue that here. 5 The existence of two choices for∆ is a reflection of the symmetry arising from the choice of direction around the cylinder which corresponds to (m, n, d ,∆) → (m, n, 1− d ,−∆) and (m, n, d ,∆) → (n, m, d ,−∆), so it is possible to force at least one of m ≤ n 1 1 or |d| < or ∆ = +1 if we wish. Indeed Jean chooses to focus in the case d < , but 2 2 here we allow either choice but recognise that the resulting intervals for d are related by this symmetry. Figure 1 shows a generating opposed pair, and Figure 2 show a variety of pairs which are not. The Fundamental Theorem gives conditions for d if (m, n) are generating and opposed. We will prove it by first finding conditions for ( m, n) to be generating. 3 Finding d given (m, n) generating The previous Theorem gives only an implicit form for d . Here we find the explicit intervals for d on which |∆ | = 1. Given m, n coprime and ∆ = ±1 we want to mn(d ) find those d such that ∆ (d ) = ∆. mn If m = n but m ̸= n we must have d = and we are done, with m = n = 1, ∆ = 1. Otherwise, assume for now that m < n. Now take u, v by solving m v − nu = ∆, specie fi d uniquely for m > 1 by 0 ≤ u < m, 0 ≤ v < n, or for m = 1 by (u, v ) = (0, 1) when ∆ = 1 or (u, v ) = (1, n − 1) when ∆ = −1. To force [md ] = u and [nd ] = v we need 1 1 u − u + 2 2 L = ≤ d ≤ = R m m m m 1 1 v − v + 2 2 L = ≤ d ≤ = R n n n n respectively and d is in the intersection of the intervals (L , R ), (L , R ). Note that n n m m eg (L , R ) is centred at v /n and has width 1/n. Then n n mn(L − L ) = ∆+ (n − m) n m mn(R − R ) = ∆− (n − m) n m mn(R − L ) = ∆+ (m + n) n m mn(R − L ) = −∆+ (m + n) m n So L > L ⇔ n > m − 2∆, which is always true unless ∆ = −1 and n = m + 1. n m Similarly R > R iff n > 2∆+ m which is true unless ∆ = 1 and n = m + 1. So apart m n from those two cases we have L < L < R < R and the interval (L , R ) is the one m n n m n n we want. To pay attention to the special cases we see that for n = m + 1 sign(L − L ) = sign(∆+ ) = ∆ n m sign(R − R ) = sign(∆− ) = ∆ n m sign(R − L ) = sign(∆+ n + ) = +1 n m sign(R − L ) = sign(−∆+ n + ) = +1 m n 6 54 54 ● ● 53 53 ● ● 52 52 ● ● 51 51 ● ● 50 50 ● ● 49 49 ● ● 48 48 ● ● 47 47 ● ● 46 46 ● ● 45 45 ● ● 44 44 ● ● 43 43 ● ● 42 42 ● ● 41 41 ● ● 40 40 ● ● 39 39 ● ● 38 38 ● ● 37 37 ● ● 36 36 ● ● 35 35 ● ● 34 34 ● ● 33 33 ● ● 32 32 ● ● 31 31 ● ● 30 30 ● ● 29 29 ● ● 28 28 ● ● 27 27 ● ● 26 26 ● ● 25 25 ● ● 24 24 ● ● 23 23 ● ● 22 22 ● ● 21 21 ● ● 20 20 ● ● 19 19 ● ● 18 18 ● ● 17 17 ● ● 16 16 ● ● 15 15 ● ● 14 14 ● ● 13 13 ● ● 12 12 ● ● 11 11 ● ● 10 10 ● ● 9 9 ● ● 8 8 ● ● 7 7 ● ● 6 6 ● ● 5 5 ● ● 4 4 ● ● 3 3 ● ● 2 2 ● ● 1 1 ● ● 0 0 (a) (5,9) is a generating but not opposed pair. (b) (9,19) is a nonopposed nongenerating pair which is not collinear. 54 54 ● ● 53 53 ● ● 52 52 ● ● 51 51 ● ● 50 50 ● ● 49 49 ● ● 48 48 ● ● 47 47 ● ● 46 46 ● ● 45 45 ● ● 44 44 ● ● 43 43 ● ● 42 42 ● ● 41 41 ● ● 40 40 ● ● 39 39 ● ● 38 38 ● ● 37 37 ● ● 36 36 ● ● 35 35 ● ● 34 34 ● ● 33 33 ● ● 32 32 ● ● 31 31 ● ● 30 30 ● ● 29 29 ● ● 28 28 ● ● 27 27 ● ● 26 26 ● ● 25 25 ● ● 24 24 ● ● 23 23 ● ● 22 22 ● ● 21 21 ● ● 20 20 ● ● 19 19 ● ● 18 18 ● ● 17 17 ● ● 16 16 ● ● 15 15 ● ● 14 14 ● ● 13 13 ● ● 12 12 ● ● 11 11 ● ● 10 10 ● ● 9 9 ● ● 8 8 ● ● 7 7 ● ● 6 6 ● ● 5 5 ● ● 4 4 ● ● 3 3 ● ● 2 2 ● ● 1 1 ● ● 0 0 (c) (5,7) is an opposed but not generating pair. (d) (1,2) is a nonopposed nongenerating pair which is collinear. Figure 2: Different types of parastichy pair in the lattice with divergence d = 17/72. 7 ∆ = 1 ∆ = −1 n ≤ m − 2 (L , R ) (L , R ) m m m m n = m − 1 (L , R ) (L , R ) n m m n n = m(= 1) n = m + 1 (L , R ) (L , R ) n m m n n ≥ m + 2 (L , R ) (L , R ) n n n n Table 1: Intervals on which (m, n) is generating in d . so if ∆ = 1 we have L < L < R < R while if ∆ = −1 it is L < L < R < R . m n m n n m n m We originally assumed m < n. If instead m > n, we can swap m and n which will change the sign of ∆, so we can summarise in Theorem 2. (m, n) is a generating pair in the lattice d iff d is in the intervals specified in Table 1. Armed with Theorem 2 we can now add the additional condition that the paras- tichy pair be opposed in order to find the Fundamental Theorem. 4 The Fundamental Theorem of Phyllotaxis 4.1 The Jean formulation Jean’s version [5, 6] of the FTP states Let (m, n) be a parastichy pair, where m and n are relatively prime, in a system with divergence angle d . The following properties are equivalent: (1) There exist unique integers 0 ≤ v < n, amd 0 ≤ u < m such that |m v − nu| = 1 and d < is in the closed interval whose end points are u/m and v /n; (2) The parastichy pair (m, n) is visible and opposed 4.2 Counterexample Consider the lattice with d = 1/12. Then the parastichy pair (m, n) = (1, 3) is neither generating nor opposed, so (2) is false. However the unique integers satisfying |m v− nu| = 1 and 0 ≤ u < 1 and 0 ≤ v < 3 are u = 0 and v = 1, and d = 1/12 < is in the interval [0/1, 1/3], so (1) is true. See Figure 3. In fact the counterexample holds for all pairs of the form (1, n), and in fairness to Jean it might be argued that 1 and n might not considered to be coprime, in which case the Theorem still holds, but this interpretation is ruled out by the comment in Appendix 4.1 that non coprime pairs are those that do not produce ‘one genetic spiral’. A more powerful defense of the utility of the Jean version of the Theorem is that it is intended for pattern recognition, typically on specimens with large, usually Fibonacci, parastichy numbers in which it is only exceptionally the case that m = 1. But even discounting the difficulty this error in the special case can cause the reader 8 54 Figure 3: (1,3) are not a generating or an opposed pair for the lattice d = 1/12. 9 in following the argument, it turns out that all modern discussions of the appearance of Fibonacci structure [8, 7, 3] invoke a successive sequence of bifurcations from more simple starting conditions, specifically (1, 1) and (1, 2), so it is important to account properly for this case. 5 Opposed generating pairs We now need to reprove the FTP, which we do by considering on which portion of the d interval where (m, n) is generating it is also opposing. First we assume ∆ > 0. d passes through 0 at L , u/m, and R and nowhere else in the interval (L , R ), m m m m m where u is defined in the proof of Theorem 2. For m > 2, by Theorem 2 the generat- ing interval is (L , R ) and we saw L < L < u/m < v /n < R < R , and so within n n m n n m the generating interval d is negative only for d < u/m and d is negative only for m n d < v /n. Thus the only region of the generating interval on which (m, n) is opposed is u/m < d < v /n. Under the symmetry, we see the analogous case for ∆ = −1. The point of this paper, though, is to define the necessary interval when m = 1. For n = 1 we have already seen we must take d = . Otherwise Theorem 2 shows the generating interval (for ∆ = +1) is of the form (1/4, 1/2) for n = 2 and 1/n ± 1/2n for n > 2. Since d changes sign every 1/2n, in either case, the generating and opposed interval for d is (1/2n, 1/n). So we can summarise in Theorem 3. ( The Fundamental Theorem of Phyllotaxis). The following are equiva- lent 1. (m ≤ n, n) is generating and opposed in the lattice d , with ∆ (d ) = ∆ mn 2. (a) m = 1, n = 1, d = , and ∆ = 1, or (b) m = 1, n > 1, ∆ = +1, d ∈ (1/2n, 1/n), or (c) m = 1, n > 1, ∆ = −1, d ∈ 1− (1/2n, 1/n) , or (d) 1 < m < n, d ∈ (u/m, v /n), ∆ = ±1, where u, v are the unique integers 0 ≤ v < n, and 0 ≤ u < m such that m v − nu = ∆. Part of the significance of this theorem, as Adler [1] and Turing [12] showed, is that if m = F and n = F are successive members of the Fibonacci sequence, then k k+1 −2 the interval for d is (F /F , F /F ) which rapidly converges to the point d = τ k−2 k k−1 k where τ is the golden ratio. 6 Discussion This correction to the Fundamental Theorem of Phyllotaxis does not reduce its cen- trality in the relationship between observed parastichy counts and the underlying mathematical structure of cylindrical lattices. Nor does recasting it partly in the ear- lier work of Turing remove the justifiable priority claims of Jean and Adler in its de- velopment, since that earlier work was languishing unpublished and incomplete in 10 the Turing archive when they independently published theirs. Nevertheless, this pa- per has taken advantage of the correction needed to the special case when one of the parastichy numbers is 1 in order to put the Theorem in a more accurate historical context and point out the common ideas of these authors. 7 Acknowledgement I’m grateful to Paul Glendinning for comments on this draft. References [1] Irving Adler. An application of the contact pressure model of phyllotaxis to the close packing of spheres around a cylinder in biological fine structure. Journal of Theoretical Biology, 67(3):447–458, August 1977. [2] P. Atela. The Geometric and Dynamic Essence of Phyllotaxis. Mathematical Modelling of Natural Phenomena, 6(02):173–186, 2011. [3] P. Atela, C. Golé, and S. Hotton. A Dynamical System for Plant Pattern Forma- tion: A Rigorous Analysis. Journal of Nonlinear Science, 12(6):641–676, March [4] S. Douady and Y. Couder. Phyllotaxis as a physical self-organized growth pro- cess. Physical Review Letters, 68(13):2098–2101, March 1992. [5] R. Jean. Number-theoretic properties of two-dimensional lattices. Journal of Number Theory, 29(2):206–223, June 1988. [6] Roger V. Jean. Phyllotaxis: A Systemic Study in Plant Morphogenesis (Cambridge Studies in Mathematics). Cambridge University Press, January 1994. [7] M. Kunz. Some analytical results about two physical models of phyllotaxis. Communications in Mathematical Physics, 169(2):261–295, May 1995. [8] G. J. Mitchison. Phyllotaxis and the Fibonacci Series. Science, 196(4287):270– 275, April 1977. [9] F. J. Richards. Phyllotaxis: Its Quantitative Expression and Relation to Growth in the Apex. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 235(629):509–564, October 1951. [10] Richard S. Smith, Soazig Guyomarc’h, Therese Mandel, Didier Reinhardt, Cris Kuhlemeier, and Przemyslaw Prusinkiewicz. A plausible model of phyllotaxis. Proceedings of the National Academy of Sciences of the United States of America, 103(5):1301–1306, January 2006. [11] Jonathan Swinton. Watching the daisies grow: Turing and Fibonacci phyllotaxis. Springer, second edition, 2004. 11 [12] Alan M. Turing. Morphogenesis, volume 3. Elsevier, 1992. http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png Quantitative Biology arXiv (Cornell University)

The Fundamental Theorem of Phyllotaxis revisited

Swinton, Jonathan
Quantitative Biology , Volume 2024 (1201) – Jan 8, 2012
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Abstract

( July 2024): This paper has now been superseded and, whilst still mathemati- cally correct, should be considered obsolete. Jean’s ‘Fundamental Theorem of Phyllotaxis’ (Phyllotaxis: a systematic study in Plant Morphogenesis, CUP 1994) describes the relationship between the count numbers of observed spirals in cylindrical lattices and the horizontal angle be- tween vertically successive spots in the lattice. It is indeed fundamental to ob- servational studies of phyllotactic counts, and especially to the evaluation of hy- potheses about the origin of Fibonacci structure within lattices. Unfortunately the textbook version of the theorem is incomplete in that it is incorrect for an important special case. This paper provides a complete statement and proof of the Theorem. Note This paper was first uploaded to arXiv in 2012, and the counterexample here to Jean’s theorem as he stated it remains valid. Since that time Infang Publishing has pub- lished my book-length treatment of Mathematical Phyllotaxis, ISBN 0993178960. This book puts the (minor) signican fi ce of the problem in more context and su- persedes this paper, which should be considered obsolete – Jonathan Swinton, July 1 Introduction Mathematical phyllotaxis is the study of the patterns that appear in two-dimensional cylindrical lattices, given particular motivation by the striking appearance of high Fibonacci numbers in a range of biological settings such as the spirals on a sun- flower [6]. Although static analyses of lattices cannot in themselves explain the ap- pearance of these numbers [12], they are essential both in relating what can actually arXiv:1201.1641v3 [q-bio.TO] 9 Jul 2024 be biologically observed to hypothesises about the underlying order, and in forming a basis for dynamical models of lattice formation [8, 4, 10, 2] that can, it is claimed, explain Fibonacci numbers and related structure in biological form. More specifi- cally, a phyllotactic theory of lattices creates a model for which lines in the lattice are most likely to be remarked on by a human observer. In the case of the sunflower or the fir cone, these lines may be those which join adjacent points in the lattice, which may be defined in different ways as contact parastichies [9, 6] or principal paras- tichies [12]. A slightly more general idea is to identify those pairs of lines that wind in opposite directions as opposed parastichies, or alternatively those lines which can be thought of as characterising the lattice, which were defined as generating parastichies by Turing [12] or equvalently as visible parastichies by Jean [6]. Jean presents the most complete description in the literature of mathematical phyllotaxis in his textbook [6], and deserves considerable credit both for innova- tion and a substantial work of integration, bringing together a range of biological datasets and historical mathematical approaches. One major contribution is what he calls the Fundamental Theorem of Phyllotaxis which he attributes in a special case to Adler [1]. The basic idea of the theorem is a very useful conceptual one. Cylindrical lattices can be, for these purposes, completely characterised by the an- gle of rotation between successive points called the divergence. If a lattice is seen to posses a specic fi pair of generating and opposed parastichies, characterised in a natural way by a pair of integers, then there is a constraint on the allowable values of the divergence. The theorem shows that for any pair of integers not both equal to 1 there are exactly two intervals of nonzero width on which the divergence will create the required generating and opposed parastichies. Unfortunately, however, in the form stated by Jean the Theorem needs modification in a range of special but important cases. The goal of this note is to restate the Theorem completely. First we give the necessary background about two-dimensional cylindrical lattices, and give a characterisation of which divergence values correspond to a given generating pair. After restating the Fundamental Theorem in Jean’s formulation and demonstrating a counter example, we then reprove a suitably corrected Theorem. Unbeknown to Jean and Adler, Turing had also considered very similar problems, but this work was unpublished at his death in 1954 [11]. It remained accessible but obscure in the Turing Archive in King’s College Cambridge, until being published in his collected works in 1992 [12], well after the relevant papers of Jean and Adler. As a secondary aim, this paper points out the ways in which Turing anticipated the later, more widely known work. 2 Background This section contains a number of statements without proof that are fairly obvi- ous on examining a diagram. They can be made rigorous by eg the use of congru- ences [12]. We consider a cylinder of circumference 1 and extending infinitely in the vertical direction, with an origin and coordinates (x , z ), 0 ≤ x ≤ 1. For any 0 ≤ d ≤ 1, we can construct a lattice d by rotating by an angle 2πd around the cylinder from the origin 2 and rising by z = 1, and repeating. This creates the set of points (x , z ) = (d , m) where m is any integer and d = md − [md ] and [x ] is the nearest integer to x , so 1 1 1 1 that − ≤ d ≤ . Since taking [ ] = 0 and [ ] = 1 map to the same point on the 2 2 2 2 cylinder we will allow the function [x ] to take the multiple values 0, 1 at the point x = . We call (d , m) the point ℓ , and from now on assume m ̸= 0. The vertical m m component is called the rise. By construction we have excluded lattices with more than one point at each rise. More generally if there are J such points spaced equally around the cylinder we would describe the lattice as having Jugacy J , but we restrict to J = 1 here. A parastichy of order m is the infinite line through the origin and ℓ with slope d /m. There are two possible lines on the cylinder through 0 and ℓ corresponding m m to winding in opposite directions and this choice of slope is equivalent to choosing the line that traverses the smallest x distance between 0 and ℓ , leaving an ambi- guity when [md ] = . The portion of this line between 0 and m defines the vector m, again with an an ambiguity when [md ] = . An m-parastichy is a member of the family of m distinct lines containing the origin-parastichy of order m and the paral- lel lines to it through the points 1, . . . , m − 1. If a point ℓ is on an m-parastichy then so is ℓ . See Figure 1. p+m A pair of (parastichy) numbers (m, n) define a pair of points ℓ , ℓ and vectors m n m,n. (m, n) is opposed if d /m and d /n have opposite sign. In the case when m n [md ] = we define the parastichy pair ( m, m) as the pair combining each of the choices of direction around the cylinder. There remains an ambiguity when [md ] = 1 1 or [nd ] = and m ̸= n which could be resolved by a specific choice of direction 2 2 although it is not of significance subsequently. There is a natural relationship between a cylindrical lattice and a corresponding periodic lattice in the plane, and the m-parastichies also dene fi an infinite family of m-parastichies in the plane lattice. A pair is generating for d if it generates the lattice in the plane in the sense that every point can be expressed as a vector sum vm+ un for integer u, v . It is necessary, but not sufficient, for a generating pair (m, n) to be coprime for if they have a common factor k all rises, including 1 must be a multiple of k . This is effectively the definition given by Turing [12], and identical to the visible pair dened fi by Jean [5, 6] in a number of different equivalent ways. Since Jean [6] gives no proof of that equivalence we give it in the Theorem below which also establishes the identity with the Turing definition, and in the process modify some of Jean’s denitions fi for extra precision. I have chosed to stick with Turing’s word generating over Jean’s visible for these identical concepts as I think the latter word carries confusing connotations in the identification of parastichy counts. We make use of the determinant ∆ of a pair (m, n), defined as mn ∆ (d ) = [nd ] m − [md ] n mn = (nd − d )m − (md − d )n n m = nd − md , m n Note that the denition fi is such because any non collinear vectors would generate the lattice in the cylinder. 3 54 Figure 1: (4,5) is a generating and opposed pair for the cylindrical d = 17/72 lattice. The parallelogram defined by the pair tiles the lattice and every lattice point is at a vertex of one of the parallelograms; the edges of the parallelograms form the paras- tichy lines. The blue lines highlight the family of 4-parastichies and the red lines the family of 5-parastichies. 4 1 1 except for the special case ∆ ( ) = m (where we have picked [ ] = 1 in the [nd ] mm 2 2 and [ ] = 0 in the [md ]). Theorem 1. (Compare Theorem 4.2 of Jean [6].) The following are equivalent 1. The pair (m, n) is generating in the lattice d . 2. The pair (m, n) has a point of the lattice d at every intersection of the lines of the pair. 3. The points 0, ℓ , and ℓ form a nondegenerate triangle which contains no m n other point of the lattice d internally. 4. (m, n) satisfy |∆ (d )| = 1. mn Proof. If a pair of vectors are collinear in the plane, they cannot be generating. If they are not collinear, the parallelogram formed by any pair can be used to tile the cylinder. This tiling will contain lattice points exactly at the vertices of each parallel- ogram iff the pair is generating, because if it has an internal point it must be a nonin- teger sum of the pair. This shows 1⇔3. Moreover the tiling produces the parastichy families of order m and n, so these must always intersect at a lattice point iff the pair is generating. This shows 1⇔2. A pair is generating iff it can express the unit vector (0 ≤ d < 1, 1) as a sum of m and n in the plane. In plane coordinates, we have [nd ]m− [md ]n = ([nd ] d − [md ] d , [nd ] m − [md ] n) m n = ([nd ] (md − [md ])− [md ] (nd − [nd ]) ,∆ ) mn = ∆ (d , 1) mn If ∆ = 1 we are done, and if ∆ = −1 we take the combination [md ]n− [nd ]m of mn mn opposite sign, so if |∆ | = 1 then (m, n) is generating. This shows 4 =⇒ 1. mn To prove 1 =⇒ 4, the central idea (of Jean and Adler and Turing) is to continue the m and n parastichies away from the origin until they cross again, so first we have to dispose of the case when the two parastichies are parallel. If m and n are paral- lel, then they are not generating, and moreover d /m = d /n so ∆ = 0 and con- m n mn versely. If they are not parallel then in the plane the parastichy of order m through (0, 0) and (d , m) and and the parastichy of order n through (1, 0) and (d , n) must m n meet at the point µ ¶ md nd mn n m = − 1, . ∆ ∆ ∆ mn mn mn If (m, n) is generating this must be a point of the lattice and so have rise equal to ′ ′ ′ both k m and k n for integer k , k , so n = k∆ and m = k ∆ . But since m and n mn mn are coprime, |∆ | = 1. mn There are close connections with the theory of Farey sequences, as mentioned in Jean [6] and in more detail in Jean [5], which can be exploited to give different versions of this proof, but we do not pursue that here. 5 The existence of two choices for∆ is a reflection of the symmetry arising from the choice of direction around the cylinder which corresponds to (m, n, d ,∆) → (m, n, 1− d ,−∆) and (m, n, d ,∆) → (n, m, d ,−∆), so it is possible to force at least one of m ≤ n 1 1 or |d| < or ∆ = +1 if we wish. Indeed Jean chooses to focus in the case d < , but 2 2 here we allow either choice but recognise that the resulting intervals for d are related by this symmetry. Figure 1 shows a generating opposed pair, and Figure 2 show a variety of pairs which are not. The Fundamental Theorem gives conditions for d if (m, n) are generating and opposed. We will prove it by first finding conditions for ( m, n) to be generating. 3 Finding d given (m, n) generating The previous Theorem gives only an implicit form for d . Here we find the explicit intervals for d on which |∆ | = 1. Given m, n coprime and ∆ = ±1 we want to mn(d ) find those d such that ∆ (d ) = ∆. mn If m = n but m ̸= n we must have d = and we are done, with m = n = 1, ∆ = 1. Otherwise, assume for now that m < n. Now take u, v by solving m v − nu = ∆, specie fi d uniquely for m > 1 by 0 ≤ u < m, 0 ≤ v < n, or for m = 1 by (u, v ) = (0, 1) when ∆ = 1 or (u, v ) = (1, n − 1) when ∆ = −1. To force [md ] = u and [nd ] = v we need 1 1 u − u + 2 2 L = ≤ d ≤ = R m m m m 1 1 v − v + 2 2 L = ≤ d ≤ = R n n n n respectively and d is in the intersection of the intervals (L , R ), (L , R ). Note that n n m m eg (L , R ) is centred at v /n and has width 1/n. Then n n mn(L − L ) = ∆+ (n − m) n m mn(R − R ) = ∆− (n − m) n m mn(R − L ) = ∆+ (m + n) n m mn(R − L ) = −∆+ (m + n) m n So L > L ⇔ n > m − 2∆, which is always true unless ∆ = −1 and n = m + 1. n m Similarly R > R iff n > 2∆+ m which is true unless ∆ = 1 and n = m + 1. So apart m n from those two cases we have L < L < R < R and the interval (L , R ) is the one m n n m n n we want. To pay attention to the special cases we see that for n = m + 1 sign(L − L ) = sign(∆+ ) = ∆ n m sign(R − R ) = sign(∆− ) = ∆ n m sign(R − L ) = sign(∆+ n + ) = +1 n m sign(R − L ) = sign(−∆+ n + ) = +1 m n 6 54 54 ● ● 53 53 ● ● 52 52 ● ● 51 51 ● ● 50 50 ● ● 49 49 ● ● 48 48 ● ● 47 47 ● ● 46 46 ● ● 45 45 ● ● 44 44 ● ● 43 43 ● ● 42 42 ● ● 41 41 ● ● 40 40 ● ● 39 39 ● ● 38 38 ● ● 37 37 ● ● 36 36 ● ● 35 35 ● ● 34 34 ● ● 33 33 ● ● 32 32 ● ● 31 31 ● ● 30 30 ● ● 29 29 ● ● 28 28 ● ● 27 27 ● ● 26 26 ● ● 25 25 ● ● 24 24 ● ● 23 23 ● ● 22 22 ● ● 21 21 ● ● 20 20 ● ● 19 19 ● ● 18 18 ● ● 17 17 ● ● 16 16 ● ● 15 15 ● ● 14 14 ● ● 13 13 ● ● 12 12 ● ● 11 11 ● ● 10 10 ● ● 9 9 ● ● 8 8 ● ● 7 7 ● ● 6 6 ● ● 5 5 ● ● 4 4 ● ● 3 3 ● ● 2 2 ● ● 1 1 ● ● 0 0 (a) (5,9) is a generating but not opposed pair. (b) (9,19) is a nonopposed nongenerating pair which is not collinear. 54 54 ● ● 53 53 ● ● 52 52 ● ● 51 51 ● ● 50 50 ● ● 49 49 ● ● 48 48 ● ● 47 47 ● ● 46 46 ● ● 45 45 ● ● 44 44 ● ● 43 43 ● ● 42 42 ● ● 41 41 ● ● 40 40 ● ● 39 39 ● ● 38 38 ● ● 37 37 ● ● 36 36 ● ● 35 35 ● ● 34 34 ● ● 33 33 ● ● 32 32 ● ● 31 31 ● ● 30 30 ● ● 29 29 ● ● 28 28 ● ● 27 27 ● ● 26 26 ● ● 25 25 ● ● 24 24 ● ● 23 23 ● ● 22 22 ● ● 21 21 ● ● 20 20 ● ● 19 19 ● ● 18 18 ● ● 17 17 ● ● 16 16 ● ● 15 15 ● ● 14 14 ● ● 13 13 ● ● 12 12 ● ● 11 11 ● ● 10 10 ● ● 9 9 ● ● 8 8 ● ● 7 7 ● ● 6 6 ● ● 5 5 ● ● 4 4 ● ● 3 3 ● ● 2 2 ● ● 1 1 ● ● 0 0 (c) (5,7) is an opposed but not generating pair. (d) (1,2) is a nonopposed nongenerating pair which is collinear. Figure 2: Different types of parastichy pair in the lattice with divergence d = 17/72. 7 ∆ = 1 ∆ = −1 n ≤ m − 2 (L , R ) (L , R ) m m m m n = m − 1 (L , R ) (L , R ) n m m n n = m(= 1) n = m + 1 (L , R ) (L , R ) n m m n n ≥ m + 2 (L , R ) (L , R ) n n n n Table 1: Intervals on which (m, n) is generating in d . so if ∆ = 1 we have L < L < R < R while if ∆ = −1 it is L < L < R < R . m n m n n m n m We originally assumed m < n. If instead m > n, we can swap m and n which will change the sign of ∆, so we can summarise in Theorem 2. (m, n) is a generating pair in the lattice d iff d is in the intervals specified in Table 1. Armed with Theorem 2 we can now add the additional condition that the paras- tichy pair be opposed in order to find the Fundamental Theorem. 4 The Fundamental Theorem of Phyllotaxis 4.1 The Jean formulation Jean’s version [5, 6] of the FTP states Let (m, n) be a parastichy pair, where m and n are relatively prime, in a system with divergence angle d . The following properties are equivalent: (1) There exist unique integers 0 ≤ v < n, amd 0 ≤ u < m such that |m v − nu| = 1 and d < is in the closed interval whose end points are u/m and v /n; (2) The parastichy pair (m, n) is visible and opposed 4.2 Counterexample Consider the lattice with d = 1/12. Then the parastichy pair (m, n) = (1, 3) is neither generating nor opposed, so (2) is false. However the unique integers satisfying |m v− nu| = 1 and 0 ≤ u < 1 and 0 ≤ v < 3 are u = 0 and v = 1, and d = 1/12 < is in the interval [0/1, 1/3], so (1) is true. See Figure 3. In fact the counterexample holds for all pairs of the form (1, n), and in fairness to Jean it might be argued that 1 and n might not considered to be coprime, in which case the Theorem still holds, but this interpretation is ruled out by the comment in Appendix 4.1 that non coprime pairs are those that do not produce ‘one genetic spiral’. A more powerful defense of the utility of the Jean version of the Theorem is that it is intended for pattern recognition, typically on specimens with large, usually Fibonacci, parastichy numbers in which it is only exceptionally the case that m = 1. But even discounting the difficulty this error in the special case can cause the reader 8 54 Figure 3: (1,3) are not a generating or an opposed pair for the lattice d = 1/12. 9 in following the argument, it turns out that all modern discussions of the appearance of Fibonacci structure [8, 7, 3] invoke a successive sequence of bifurcations from more simple starting conditions, specifically (1, 1) and (1, 2), so it is important to account properly for this case. 5 Opposed generating pairs We now need to reprove the FTP, which we do by considering on which portion of the d interval where (m, n) is generating it is also opposing. First we assume ∆ > 0. d passes through 0 at L , u/m, and R and nowhere else in the interval (L , R ), m m m m m where u is defined in the proof of Theorem 2. For m > 2, by Theorem 2 the generat- ing interval is (L , R ) and we saw L < L < u/m < v /n < R < R , and so within n n m n n m the generating interval d is negative only for d < u/m and d is negative only for m n d < v /n. Thus the only region of the generating interval on which (m, n) is opposed is u/m < d < v /n. Under the symmetry, we see the analogous case for ∆ = −1. The point of this paper, though, is to define the necessary interval when m = 1. For n = 1 we have already seen we must take d = . Otherwise Theorem 2 shows the generating interval (for ∆ = +1) is of the form (1/4, 1/2) for n = 2 and 1/n ± 1/2n for n > 2. Since d changes sign every 1/2n, in either case, the generating and opposed interval for d is (1/2n, 1/n). So we can summarise in Theorem 3. ( The Fundamental Theorem of Phyllotaxis). The following are equiva- lent 1. (m ≤ n, n) is generating and opposed in the lattice d , with ∆ (d ) = ∆ mn 2. (a) m = 1, n = 1, d = , and ∆ = 1, or (b) m = 1, n > 1, ∆ = +1, d ∈ (1/2n, 1/n), or (c) m = 1, n > 1, ∆ = −1, d ∈ 1− (1/2n, 1/n) , or (d) 1 < m < n, d ∈ (u/m, v /n), ∆ = ±1, where u, v are the unique integers 0 ≤ v < n, and 0 ≤ u < m such that m v − nu = ∆. Part of the significance of this theorem, as Adler [1] and Turing [12] showed, is that if m = F and n = F are successive members of the Fibonacci sequence, then k k+1 −2 the interval for d is (F /F , F /F ) which rapidly converges to the point d = τ k−2 k k−1 k where τ is the golden ratio. 6 Discussion This correction to the Fundamental Theorem of Phyllotaxis does not reduce its cen- trality in the relationship between observed parastichy counts and the underlying mathematical structure of cylindrical lattices. Nor does recasting it partly in the ear- lier work of Turing remove the justifiable priority claims of Jean and Adler in its de- velopment, since that earlier work was languishing unpublished and incomplete in 10 the Turing archive when they independently published theirs. Nevertheless, this pa- per has taken advantage of the correction needed to the special case when one of the parastichy numbers is 1 in order to put the Theorem in a more accurate historical context and point out the common ideas of these authors. 7 Acknowledgement I’m grateful to Paul Glendinning for comments on this draft. References [1] Irving Adler. An application of the contact pressure model of phyllotaxis to the close packing of spheres around a cylinder in biological fine structure. Journal of Theoretical Biology, 67(3):447–458, August 1977. [2] P. Atela. The Geometric and Dynamic Essence of Phyllotaxis. Mathematical Modelling of Natural Phenomena, 6(02):173–186, 2011. [3] P. Atela, C. Golé, and S. Hotton. A Dynamical System for Plant Pattern Forma- tion: A Rigorous Analysis. Journal of Nonlinear Science, 12(6):641–676, March [4] S. Douady and Y. Couder. Phyllotaxis as a physical self-organized growth pro- cess. Physical Review Letters, 68(13):2098–2101, March 1992. [5] R. Jean. Number-theoretic properties of two-dimensional lattices. Journal of Number Theory, 29(2):206–223, June 1988. [6] Roger V. Jean. Phyllotaxis: A Systemic Study in Plant Morphogenesis (Cambridge Studies in Mathematics). Cambridge University Press, January 1994. [7] M. Kunz. Some analytical results about two physical models of phyllotaxis. Communications in Mathematical Physics, 169(2):261–295, May 1995. [8] G. J. Mitchison. Phyllotaxis and the Fibonacci Series. Science, 196(4287):270– 275, April 1977. [9] F. J. Richards. Phyllotaxis: Its Quantitative Expression and Relation to Growth in the Apex. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 235(629):509–564, October 1951. [10] Richard S. Smith, Soazig Guyomarc’h, Therese Mandel, Didier Reinhardt, Cris Kuhlemeier, and Przemyslaw Prusinkiewicz. A plausible model of phyllotaxis. Proceedings of the National Academy of Sciences of the United States of America, 103(5):1301–1306, January 2006. [11] Jonathan Swinton. Watching the daisies grow: Turing and Fibonacci phyllotaxis. Springer, second edition, 2004. 11 [12] Alan M. Turing. Morphogenesis, volume 3. Elsevier, 1992.

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Quantitative Biology – arXiv (Cornell University)

Published: Jan 8, 2012

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