1 Appendix A. Derivation of Length Asymmetry Ratio (ππ ) Given Branching Angles, 2 Here, we relate the length asymmetry ratio (ππ ) to the optimal branching solutions (ππ ) 3 and the geometry of the unshared endpoints (i.e., the vertices ππ ). We denote the 4 lengths of the sides of the triangle that correspond to π1 π2, π0 π2 , and π0 π1 as π£0 , π£1 , 5 and π£2 , respectively (Fig A1). We first prove two Lemmas that lead to the derivation of 6 the asymmetric length ratio. 7 Lemma 1: Let the intersection of the line between the points π0 and π½ with the line π1 π2 8 be called πΎ and the angle defined by the three points πΜ 0 πΎπ1 be called π (Fig A1). Using 9 these definitions and the other labeling in Fig. A1, the following relationships holds 10 11 12 |π1 πΎ| π1 sin π2 π£2 sin π1 = = |π2 πΎ| π2 sin π1 π£1 sin π2 Proof: By the law of sines applied to the triangles π0 π1 πΎ and π0 π2 πΎ, we have: sin π sin π1 = , π£2 |π1 πΎ| sin(π β π) sin π2 = π£1 |π2 πΎ| |π πΎ| π£ sin π 13 Since sin(π β π) = sin π, dividing these equations yields |π1 πΎ| = π£2 sin π1. Applying a 14 similar approach to triangles π½π1 πΎ and π½π2 πΎ, we have |π1 πΎ| = π1 sin π2 , as desired. β 15 Figure A1. (a) Schematic of the branching geometry (b) Illustration of degenerate cases 16 where the branching junction coincides with one of the vertices. 2 |π πΎ| 2 1 2 π sin π 2 1 17 π 18 Lemma 2: The length asymmetry ratio (ππ = π1) can be calculated purely in terms of the 19 lengths of the sides of π£1 and π£2 along with the angle π1Μ π0 π2 and the branching 20 angles π1 and π2 as 2 ππ = 21 π£2 sin π1 (β cos π1Μ π0 π2 + sin π1Μ π0 π2 cot(π1Μ π0 π2 + πΎ β π2 )) π£1 sin π2 π£2 sin π1 +cos(π1 +π2 βπ1Μ π0 π2 ) β1 π£1 sin π2 cot [ ] Μ sin(π1 +π2 βπ1 π0 π2 ) 22 where πΎ = 23 Proof: By Lemma 1, we have ππ = 24 π1 π£2 sin π1 sin π1 = π2 π£1 sin π2 sin π2 25 Then, by applying law of sines in a specific, successive order and also using sine 26 addition formulas, we express sin π1 in terms of known quantities and branching angles: 27 sin π1 = (β cos π1Μ π0 π2 + sin π1Μ π0 π2 cot(π1Μ π0 π2 + πΎ β π2 )) sin π2 28 sin π 2 proving the lemma. β 29 With Lemma 2, we show that the branching angle solutionβobtained by optimizing 30 certain structural principlesβalso predicts the optimal value for the asymmetric length 31 ratio. 32 Appendix B. Coordinate-Free Framework for Material Cost Optimization Solutions 33 In this section, we introduce a coordinate-free framework for the minimization of the 34 objective function, defined as π» = βπ βπ ππ . We have not seen this approach in the 35 literature, and other references have used methods that rely on specific choices of 36 coordinate systems and complicated algebra (1-3). The solution is obtained via finding 37 the stationary and singular points of the cost function π» with respect to π0 (the parent 38 vessel length) and π1 (the angle of the parent vessel relative to its unshared 39 endpoint π0) (Fig A1). Below, we provide two lemmas that will be used to determine ππ 40 and ππ . 41 Lemma 3. Given fixed endpoints π0 , π1 , and π2, the length |π0 π1 | and the angle π1 are 42 fixed in the triangle π0 π½π1, (Fig A2), the derivative of a daughter vessel length with 43 respect to the parent vessel length is ππ» 0 ππ» 1 44 ππ1 = cos π2 ππ0 45 Proof: Draw a perpendicular line passing through π1 and intersecting with the extension 46 of π0 π½ at π. Denote |π0 π1 | = π£2 , |π1 π| = π¦, and |π½π| = π₯. When J is on the right side of 47 V0, we have π£2 cos π = π₯ + π0 . Since π£2 cos π1 is fixed because π£2 and π1 are fixed, it 48 follows that π(π£2 cos π) = π(π₯ + π0 ) = 0, or equivalently ππ₯ = β1. ππ0 (A1) ππ₯ 49 Notice however that the derivative ππ is discontinuous when the branching junction 50 collapses on the parent endpoint (i.e., π0 = 0) as the right and left derivatives of π₯ with 51 respect to π0 are opposite in sign: π+ π₯(0) = 52 1 (Fig A2). 53 Figure A2. (a) The branching geometry of a parent and one of the daughter vessels (b) 54 When the vertex π½ approaches the vertex π0 from the right, π₯ = π£2 πππ π1 β π0. (c) When 55 the vertex π½ approaches the vertex V0 from the right, π₯ = π£2 πππ π1 + π0. 0 π(π£2 cos π1 βπ0 ) ππ0 = β1, πβ π₯(0) = π(π£2 cos π1 +π0 ) ππ0 = 56 57 Applying the Pythagorean Theorem to the triangle π1 π½π, we have π1 = βπ₯ 2 + π¦ 2 , hence ππ1 π₯ π₯ = = ππ₯ βπ₯ 2 + π¦ 2 π1 58 (A2) Using the chain rule along with equations (A1) and (A2) gives ππ1 ππ1 ππ₯ ππ1 π₯ = =β = β = β cos(π β π2 ) = cos π2 ππ0 ππ₯ ππ0 ππ₯ π1 59 60 as desired. 61 Lemma 4. Given fixed lengths |π0 π1 | = π£2 and π0 in the triangle π0 π½π1, then 62 ππ1 = βπ0 sin π2 ππ1 63 Proof: As in Lemma 1, we have cos π2 = βcos(π β π2 ) = β π and π1 = βπ₯ 2 + π¦ 2 . From 64 the triangle π0 π1 π, we further have π¦ = π£2 sin π1 and π₯ = π£2 cos π1 β π0 . Substituting 65 these into the expression for π1 yields π1 = β(π£2 cos π1 β π0 )2 + (π£2 sin π1 )2. As π£2 and π0 66 are fixed, differentiating π1 with respect to π1 gives: π₯ 1 67 ππ1 1 2(π£2 cos π1 β π0 )(βπ£2 sin π1 ) + 2π£22 sin π1 cos π1 = ππ1 2 β(π£2 cos π1 β π0 )2 + (π£2 sin π1 )2 68 This expression simplifies by cancelling the 2π£22 sin π1 cos π1 terms in the numerator and 69 by recognizing the denominator is equal to π1. Therefore, we obtain 70 Since sin π1 = π£ and sin(π β π2 ) = π , this equation becomes π¦ π¦ 2 1 ππ1 ππ1 = π0 π£2 π1 sin π1 . ππ1 π0 π£2 π¦ = sin π = π0 = βπ0 sin π2 β ππ1 π1 π1 71 72 With these two lemmas proven, we now return to the original optimization problem. 73 Unless J coincides with the unshared endpoints π0 , π1 or π2 , substituting Lemma 1 and 74 Lemma 2 into the equality, we have ππ» ππ» βπ» = ( , ) = ββ0, ππ0 ππ1 75 76 leads to two equations β0 = ββ1 cos π2 β β2 cos π1 (A3) β1 π ππ π2 = β2 π ππ π1 (A4) 77 Solving these equations yields the previously reported branching angle solutions (Eq. 78 (1) in our paper and from Zamir et. al. (1, 2)). Dividing both sides of the equations (A3) and (A4) by β2 and combining them, we 79 80 81 have β0 sin π1 πππ π2 + sin π2 πππ π1 β sin(π1 + π2 ) =β = β2 sin π2 sin π2 82 Realizing that π1 + π2 = 2π β π0 , or equivalently β sin(π1 + π2 ) = sin π0 , and combining 83 the above equations (A3) and (A4) yields β0 π ππ π2 = β2 π ππ π0 . Thus, in order for the 84 equations that follow from βπ» = β0 to be soluble, the expressions sin π0 , π ππ π1 , and sin π2 85 must all have the same sign because the length scales hi are all positive. This sign 86 criterion can only be satisfied when the branching junction is inside of the triangle 87 defined by π0 , π1 and π2. Consequently, this implies βπ» = β0 cannot be satisfied when the 88 branching junction is outside of the triangle or on the boundary of the triangle. 89 Therefore, in order for the previously reported formula for the branching-angle solutions 90 to be valid, we need to check first if β1 β€ cos ππ β€ 1, and if it does not, we must 91 conclude that βπ» = β0 does not have a solution. Previous studies were not explicit about 92 this criterion or distinction in finding solutions. Solving the inequalities β1 β€ cos ππ β€ 1 93 for each branching angle yields necessary conditions for the existence of solutions 94 of βπ» = β0. These conditions reduce to the simple statement, βπ < βπ + βπ , about the 95 weightings of the terms in the cost function for any combination of (π, π, π). If any of these 96 three conditions fail, then the branching junction will be degenerate, meaning that the 97 optimal branching junction, J, will collapse to one of the vertices. 98 Moreover, the angles of the triangle π0 π1 π2 further confine the range of branching 99 angles that can be realized within the triangle, i.e. πΜ π ππ ππ < ππ . Hence, if branching angle 100 solutions defined by Eq. (1) violate any of these conditions, the optimization solution will 101 be a collapse of the branching junction onto one of the unshared endpoints. 102 Appendix C. Degeneracy Solutions of Material Cost Optimization 103 We now derive which particular vertex the branching junction will collapse onto for the 104 degeneracy cases. 105 Lemma 5. When the triangle conditions and inequalities do not hold (i.e., βπ β₯ βπ + βπ ), 106 the vertex ππ associated with the largest cost parameter (i.e., βπ ) is the solution for the 107 material cost optimization. 108 Proof: By symmetry and without loss of generality, we assume that the cost per parent 109 length is greater than the sum of the costs per length for the daughter vessels, i.e. β0 β₯ 110 β1 + β2 . To identify the vertex that yields the minimum cost, we will calculate the total 111 cost corresponding to all three degenerate cases (Fig A1). Total costs at the 112 corresponding vertices are given by π»π0 = β1 π£2 + β2 π£1 , π»π1 = β0 π£2 + β2 π£0 , and π»π2 = 113 β0 π£1 + β1 π£0 , where π£0 , π£1 , and π£2 are lengths of sides π1 π2 , π0 π2 , π0 π1 respectively. From 114 our assumption and triangle inequality applied to the sides of the triangle π0 π1 π2 , we 115 have π»π1 = β0 π£2 + β2 π£0 β₯ (β1 + β2 )π£2 + β2 π£0 = β2 (π£0 + π£2 ) + β1 π£2 > β2 π£1 + β1 π£2 = π»π0 . 116 In a symmetric way, one can also prove that π»π2 > π»π1 , implying that J collapses on π0. 117 Lemma 6. For any triangle with vertices X, Y, Z, and a point P inside this triangle we 118 have the following inequality |ππ| + |ππ| > |ππ| + |ππ| 119 120 Proof: The set of points Yβ on the plane for which |ππβ²| + |πβ²π| = |ππ| + |ππ| 121 122 forms an ellipse as illustrated in Fig A3. Therefore, for any point πβ² in the interior of the 123 ellipse |ππβ²| + |πβ²π| < |ππ| + |ππ| 124 125 proving the claim. 126 Fig A3. Ellipse formed by the points X, Y, and Z. By definition, the sum of the distances 127 from any point on the ellipse to X and Z is fixed. 128 129 Lemma 7. When optimal branching angle solutions (Eq. (1)) result in a case where the 130 triangle condition (πΜ π ππ ππ β₯ ππ ) fails, then the vertex associated with ππ for which the 131 inequality fails also provides the minimum of π». 132 Proof: Without loss of generality, let us assume that the optimal solution of π0 is less 133 than the angle π1Μ π0 π2 . As β0 2 = β1 2 + β2 2 β 2β1 β2 cos(π β π0 ), we can form a triangle 134 OAB with side-lengths β0 , β1 , β2 that has the angle π β π0 at the vertex A (Fig A4). Now, 135 let us construct a triangle ABC similar to the triangle π0 π1 π2. Drawing a line segment AC 136 Μ equals πΜ0 β π2Μ of length β2 π£1 , so that the angle πΆπ΄π΅ π0 π1 , yields such a triangle with 137 similarity ratio 138 applied to the concave quadrilateral OBCA (Lemma 6) leads to β0 +β2 π£0 > β1 +β2 π£1. 139 Multiplying both sides by π£2 provides π»π1 = β0 π£2 + β2 π£0 > β2 π£1 + β1 π£2 = π»π0 . In a similar 140 manner, we can show that π»π2 > π»π0 , demonstrating that π0 gives the optimal position 141 for J. By symmetry, when π1 < πΜ 0 π1 π2 this implies the branching junction J collapses to 142 π1, and when π2 < π1Μ π2 π0, this implies that J collapses to π2. 143 Fig A4. The diagram of the proof to show showing that when π < πΜ0 , the branching 144 junction J will collapse on π0. π£ 2 β2 π£2 π£ . Hence, the side BC has length β2 π£0 (Fig A4). Then, the side inequality 2 π£ 2 π£ 2 145 146 Appendix D. Power Cost Optimization for a Single Branching Junction Solutions 147 Here, we show that power cost optimization always leads to degenerate branching 148 geometry. To do this, we first calculate the equivalent impedances when the branching 149 junction J occurs at the vertex ππ (Fig A1)βdenoted by πππ βfor each i . 150 ππ0 1 1 β1 =( + ) , β1 π£2 β2 π£1 ππ1 = β0 π£2 , ππ2 = β0 π£1 151 Now, if we show that πππ β₯ min(ππ0 , ππ1 , ππ2 ), it follows that πππ attains its minimum at 152 one of the vertices. Without loss of generality, we assume that π£1 β€ π£2 , so ππ2 β€ ππ1 153 and min(ππ0 , ππ1 , ππ2 ) = min(ππ0 , ππ2 ). The following lemmas verify our claim that one of 154 the vertices is always optimal for the branching junction. 155 Lemma 8. Let ππ0 < ππ2 . Then, min(πππ ) = ππ0 156 Proof: To prove the lemma, we need to show that πππ β₯ ππ0 for all possible locations of 157 the branching junction, J. Because ππ0 < ππ2 , we have β0 > π£ (β 158 form the following inequality by replacing β0 by (β 159 160 161 162 1 1 π£2 1 1 1 π£2 +β 1 2 π£1 1 +β 1 2 π£1 β1 ) , so we can β1 1 ) π£1 1 1 β1 1 1 β1 π0 1 1 β1 πππ = β0 π0 + ( + ) >( + ) +( + ) β1 π1 β2 π2 β1 π£2 β2 π£1 π£1 β1 π1 β2 π2 To prove πππ β₯ ππ0 = (β 1 1 π£2 +β 1 2 π£1 β1 ) , it suffices to prove 1 1 β1 π0 1 1 β1 1 1 β1 ( + ) +( + ) β₯( + ) β1 π£2 β2 π£1 π£1 β1 π1 β2 π2 β1 π£2 β2 π£1 Rearranging terms, the proof of the Lemma boils down to proving the inequality 1 1 β1 π0 1 1 β1 ( + ) > (1 β ) ( + ) , β1 π1 β2 π2 π£1 β1 π£2 β2 π£1 (A5) 1 1 163 Taking the reciprocal of both sides of (A5) and factoring out the terms with β and β , this 164 inequality is equivalent to 1 2 1 1 1 π0 β1 1 1 1 π0 β1 ( β (1 β ) ) + ( β (1 β ) ) < 0 β1 π1 π£2 π£1 β2 π2 π£1 π£1 165 166 Hence, if we show that both of the terms in the above expression are negative, then 167 their sum would also be negative, and the proof will be complete. In other words, it 168 suffices to show two inequalities 169 170 1 1 π0 β1 β (1 β ) < 0 π1 π£2 π£1 (A6) 1 1 π0 β1 β (1 β ) < 0 π2 π£1 π£1 (A7) Observe that the triangle inequality applied to the triangle π0Jπ1 gives π0 + π1 > π£2 , hence π1 π£2 π π > 1 β π£0 > 1 β π£0 , proving (A6). Moreover, the triangle inequality applied to the triangle 2 1 π π 171 π0Jπ2 yields π0 + π2 > π£1 , implying that π£2 > 1 β π£0 , which proves (A7). β 172 The next lemma takes care of the complementary case. 173 Lemma 9: Let ππ0 > ππ2 , then min πππ = ππ2 174 Proof: Following the same idea as in the proof of Lemma 8, we want to show that πππ β₯ 175 ππ2 , or equivalently 1 1 1 1 β1 ( + ) > β0 (π£1 β π0 ) β1 π1 β2 π2 176 177 By the inequality (A5), we proved in Lemma 1, we have 1 1 β1 π0 1 1 β1 ( + ) > (1 β ) ( + ) β1 π1 β2 π2 π£1 β1 π β2 π£1 178 179 The assumption (β 1 1 π£2 +β 1 2 π£1 β1 ) > β0 π£1 further yields that 1 1 β1 π0 ( + ) > (1 β ) β0 π£1 = β0 (π£1 β π0 ) β1 π1 β2 π2 π£1 180 181 as desired. β 182 With Lemmas 8 and 9, we proved that the branching junction collapses onto one of the 183 vertices for any choice of cost parameters β0 , β1 , and β2 . 184 Appendix E. Enlarged Consideration of the Power Cost Optimization to Go 185 Beyond a Single Branching 186 In this section, we add terms π1 and π2 to the calculation of πΜππ to respectively represent 187 the impedance of all of the vessels are downstream from each daughter vessel at that 188 branching junction. Furthermore, we consider the special case that impedance 189 matchingβthe impedance of the parent vessel is matched by the equivalent 190 impedances of the daughter vesselsβis satisfied throughout the whole network. By 191 requiring that siblings have identical impedances and that each sibling has the same 192 number of downstream vessels, we show that the ratio ππ is larger for vessels that are 193 near to the first branching level (i.e., the heart). To simplify the calculations, we π π 194 enumerate the levels such that the level number increases from capillary (level 0) to the 195 heart (level N). This is the reverse of the labeling used in most models. 196 By applying impedance matching successively from level 0 to level k, we first 197 recognize that the impedance of the vessel at the πth level is given by π0 /2π , where 198 π0 denotes the impedance of the capillary. Moreover, for the first few levels above the 199 capillary level (when π = 0, 1, 2), we find that the downstream impedance at level k 200 follows the form 201 Lemma 10. The downstream impedance from a daughter vessel at level k is given by ππ 2π (Fig A5). The next Lemma generalizes this formula for all levels k. ππ = 202 ππ0 2π 203 Proof: We prove this claim by induction. Note that a vessel at level π β 1 is in series 204 with the downstream vessels as illustrated in the Fig A5. If the downstream impedance 205 at level (π β 1) is equal to 206 impedance at level k is given by 207 208 209 ππ = (πβ1)π 2πβ1 , then by rules of fluid mechanics, the downstream 1 1 1 + (π β 1)π0 (π β 1)π0 π0 π0 + + 2πβ1 2πβ1 2πβ1 2πβ1 = ππ0 .β 2π Hence, by Lemma 9, we have that the value of ππ /ππ at level π is equal to ππ0 2π = π π0 2π 210 so that the value of this ratio increases with the level (i.e., increase from the capillaries 211 to the heart). Therefore, near the heart, the constants (ππ ) representing the downstream 212 impedances in the optimization scheme are relatively large compared to the 213 impedances (ππ ) of the daughter vessels at that branching junction. 214 Figure A5. (a) Perfectly-balanced branching network with identical daughter 215 impedances and (b) inclusion of impedances for downstream vessels in entire 216 branching network and thus beyond just the branching level k. 217 218 219 References 220 1. 221 general physiology. 1978;72(6):837-45. 222 2. 223 Biology. 1976;62(1):227-51. 224 3. 225 branching of arteries. The Journal of general physiology. 1926;9(6):835-41. 226 Zamir M. Nonsymmetrical bifurcations in arterial branching. The Journal of Zamir M. Optimality principles in arterial branching. Journal of Theoretical Murray CD. The physiological principle of minimum work applied to the angle of
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