This poster was presented at the international genetics convention in Brisbane in 2006.
Effect of Size on Virtual Population Growth


Sibly et al estimated the relationship between a population's size and its growth rate for 1,780 time series of mammals, birds, fish, and insects. Growth rates are high at low population densities but, contrary to previous predictions, decline rapidly with increasing population size and then flatten out, for all four taxa. This produces a strongly concave relationship between a population's growth rate and its size.  The fact that the curve is defined by only three values (vertical and horizontal position and degree of curvature) suggests a simple scientific law lying at the very heart of evolution and, we suggest, genetics.  We undertook to duplicate the curve with one of two programs we had available that could simulate a population, specify details of its genetic behavior and follow it over generations. 

                     A                                            B

Recent work on the speciation of Anopheles mosquitoes (Microarrays and Species Origins.  Roger Butlin and Cally Roper.  NATURE 437: 199-201 SEPTEMBER 2005) has found that “speciation genes,” which appear to be causing speciation, are located near the centromeres (A) of the chromosomes, a region that does not undergo crossing over. 

Of two programs we had, one modeled genes that assorted independently or were loosely linked, which would be like genes located at a distance from the centromere (B) or on different chromosomes.  The other program had all genes rigidly bound to one pair of chromosomes.  This proved to be the program that was able to reproduce the concave curve described by Sibly et al.

Spread Sheet of Growth Rate of Different Population Sizes

max pop

 

0

20

220

420

620

820

1020

1

 

 

48

344

0

861

0

1041

2

 

 

0

313

611

885

1021

966

3

 

 

48

384

503

925

1023

1215

4

 

 

29

378

609

871

1096

1301

5

 

 

44

261

611

859

882

1073

6

 

 

36

282

568

827

0

0

7

 

 

26

238

598

930

996

175

8

 

 

53

372

435

709

0

1086

9

 

 

47

283

451

719

0

0

10

 

 

40

313

589

788

1190

0

sum

 

0

371

3168

4975

8374

6208

6857

extincts

 

 

1

0

1

0

4

3

div by 10

 

0

37.1

316.8

497.5

837.4

620.8

685.7

growth 1

 

0

17.1

96.8

77.5

217.4

-199.2

-334.3

rate 1

 

#DIV/0!

0.855

0.44

0.184524

0.350645

-0.24293

-0.32775

% no 1

 

#DIV/0!

85.5

44

18.45238

35.06452

-24.2927

-32.7745

survivors

 

10

9

10

9

10

6

7

sum by survivors

0

41.22222

316.8

552.7778

837.4

1034.667

979.5714

growth 2

 

0

21.22222

96.8

132.7778

217.4

214.6667

-40.4286

rate 2

 

#DIV/0!

1.061111

0.44

0.316138

0.350645

0.261789

-0.03964

% no 2

 

#DIV/0!

106.1111

44

31.61376

35.06452

26.17886

-3.96359

extinct gens

 

 

 

 

 

 

 

1

 

 

444

 

833

 

215

995

2

 

 

0

 

 

 

954

750

3

 

 

0

 

 

 

443

402

4

 

 

0

 

 

 

356

 

5

 

 

0

 

 

 

 

 

6

 

 

0

 

 

 

 

 

7

 

 

0

 

 

 

 

 

8

 

 

0

 

 

 

 

 

9

 

 

0

 

 

 

 

 

10

 

 

0

 

 

 

 

 

sum extinct gens

0

444

0

833

0

1968

2147

av time to extinctin

#DIV/0!

444

#DIV/0!

833

#DIV/0!

492

715.6667

loss per gen per run

#DIV/0!

0.045045

#DIV/0!

0.504202

#DIV/0!

1.666667

1.425245

total extinction loss

#DIV/0!

0.045045

#DIV/0!

0.504202

#DIV/0!

6.666667

4.275734

sum less extinctions

#DIV/0!

370.955

#DIV/0!

4974.496

#DIV/0!

6201.333

6852.724

div by 10

 

#DIV/0!

37.0955

#DIV/0!

497.4496

#DIV/0!

620.1333

685.2724

growth 3

 

#DIV/0!

17.0955

#DIV/0!

77.44958

#DIV/0!

-199.867

-334.728

rate 3

 

#DIV/0!

0.854775

#DIV/0!

0.184404

#DIV/0!

-0.24374

-0.32816

% no 3

 

#DIV/0!

85.47748

#DIV/0!

18.44038

#DIV/0!

-24.374

-32.8164


1220

1420

1620

1820

2020

1906

0

1839

0

2032

1559

662

1957

0

0

0

0

0

0

2391

1675

2158

0

0

0

495

0

0

0

2534

0

0

2038

1986

0

0

0

0

2162

0

1382

0

0

0

0

1612

1320

0

0

0

0

0

0

0

2391

8629

4140

5834

4148

9348

4

7

7

8

6

862.9

414

583.4

414.8

934.8

-357.1

-1006

-1036.6

-1405.2

-1085.2

-0.2927

-0.70845

-0.63988

-0.77209

-0.53723

-29.2705

-70.8451

-63.9877

-77.2088

-53.7228

6

3

3

2

4

1438.167

1380

1944.667

2074

2337

218.1667

-40

324.6667

254

317

0.178825

-0.02817

0.200412

0.13956

0.156931

17.88251

-2.8169

20.04115

13.95604

15.69307

 

 

 

 

 

71

349

152

135

86

802

240

379

299

883

580

308

629

946

143

884

615

64

109

68

 

756

638

211

962

 

122

68

187

331

 

186

68

765

 

 

 

 

185

 

 

 

 

 

 

 

 

 

 

 

2337

2576

1998

2837

2473

584.25

368

285.4286

354.625

412.1667

2.088147

3.858696

5.675676

5.132182

4.90093

8.352589

27.01087

39.72973

41.05746

29.40558

8620.647

4112.989

5794.27

4106.943

9318.594

862.0647

411.2989

579.427

410.6943

931.8594

-357.935

-1008.7

-1040.57

-1409.31

-1088.14

-0.29339

-0.71035

-0.64233

-0.77434

-0.53868

-29.339

-71.0353

-64.2329

-77.4344

-53.8683

The parameters for these runs were: Max generations 1000 . Max offspring 6.  Initial population 100. Max Population independent variable.  Gene pairs subject to recessive lethal mutations 100.  Mutation rate of the genes 10.  Mutually tuned components 100.  Mutation rate for the mutually tuned systems was 400.  Thousandths of offspring lost per unit of detuning was 40. 


Graph of Growth Rate against Population Size


Simply assigning 0 population to the extinct line is “% no 1” on the graph.  “% no 2,” dropping the populations that go extinct from consideration, underestimates growth reduction, but is the approach used by the authors.  They, too, observed that local populations go extinct in the wild all the time.  % no 3 takes the maximum population size, divided by the number of generations it took to go extinct, and winding up with a number lost per generation, subtracting that from the sum of the offspring in the populations that survived.  This approach is still not perfect, but does give us a line, % no 3, which is essentially the same as % no 1.  Compare with page I.

Where Evolution Places a Genome

The model accounts for observations, but there are more variables in the model than in reality.  There must be some simplifying mechanism.  That mechanism is evolution.  In the chart A, B, and C are forbidden because the excessive mutation rate would produce extinction.  E is not plausible.  D we doubt to be possible.  F is possible for a finite time, but evolution will increase complexity in the direction of arrow G, until complexity is maximized in H.  This is an intrinsic limit to evolution.  (In Search of the Limits of Evolution.  Fyodor A. Kondrashov  NATURE GENETICS: 17: 9 January, 2005) .  A subset of H, I, will have the properties we have shown.  The fact that evolution must stop in H explains punctuated equilibrium.  Evolution starts anew when a form is returned to F because of a favorable change that permits survival with less performance, such as domestication or monogamy. Driving all forms into I reduces the variability that can be observed.

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