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Liam McNulty takes you through some experiments with the Neighbour-Sensing model of mycelial growth |
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This page summarises a series of experiments that explore some of the capabilities of the Neighbour-Sensing model. The experiments were done by changing the parameters that the simulation uses in its mathematical computations, so that you can SEE on screen the effects of those parameters on growth of the fungus.
What follows in this document is a record of these experiments presented in tabulated form standardised so that in most cases the left cell shows the image of the mycelium generated by the parameter set, and the right cell lists the parameters used to produce that image. The definitions of the parameters are laid out so that they reflect the lay out of the parameters page of the program itself (Fig. 1).
These experiments used a version of the program in which the user interface for defining the parameters to be used in each run had the appearance shown in Fig. 1. Following this figure is a series of experimental records are presented so that accompanying the image of the mycelium is a textual definition of the parameters used to produce the image, these being laid out to reflect the lay out of the parameters interface page of the program (Fig. 1).
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Fig. 1. User interface for defining the parameters to be used in each run (early version of the program). |
In the printed parameter sets a hyphen represents an unchecked box in the parameter dialogue (see Fig. 1), and the parameter(s) shown in red-bold font are the ones being illustrated. The time units shown above each parameter set should be viewed as consecutive blocks during which that parameter set has been operative. Each figure has a standard scale bar of 100 length units. Simulations can be paused, parameters changed, and the simulation resumed. In these cases several parameter sets are shown accompanying a single illustration.
Figs 2-4 show the effects of varying the autotropism setting.
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100 time units: - (no negative autotropism) - branch only when less than 3 grow only when less than 15 neighbouring tips in radius = 20 - - - branching probability = 40 growth speed = 1 |
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Fig. 2. Growth pattern without negative autotropism |
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100 time units: suppose negative autotropism 0.5 - branch only when less than 3 grow only when less than 15 neighbouring tips in radius = 20 - - - branching probability = 40 growth speed = 1 |
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Fig. 3. Growth pattern with negative autotropism implemented at a value of 0.5. |
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100 time units: suppose negative autotropism 0.1 - branch only when less than 3 grow only when less than 15 neighbouring tips in radius = 20 - - - branching probability = 40 growth speed = 1 |
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Fig. 4. Growth pattern with negative autotropism set to 0.1. |
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Fig. 4 shows the growth pattern obtained with negative autotropism set to 0.1. This seems to produce the most realistic mycelial shapes and was the value used in most of the subsequent examples shown here.
Figs 5 and 6 illustrate growth patterns produced with different levels of control over the probability of branching (but with the same growth rules).
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100 time units: suppose negative autotropism 0.1 - branch only when less than 2 - neighbouring tips in radius = 20 - - - branching probability = 40 growth speed = 1 |
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Fig. 5. Growth pattern obtained by setting a low threshold for preventing branching (i.e. making branching rare) (compare with Fig. 6).
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100 time units: suppose negative autotropism 0.1 - branch only when less than 8 - neighbouring tips in radius = 20 - - - branching probability = 40 growth speed = 1 |
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Fig. 6. Increasing the threshold for preventing branching (i.e. making branching more frequent) results in a much denser structure (compare with Fig. 5). |
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Introducing a limitation on the growth of tips modifies this colony morphology further, resulting in a structure of intermediate density (Fig. 7). However, if this parameter is implemented its value must be larger than that of the branching threshold parameter for a viable, growing mycelium to result.
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100 time units: suppose negative autotropism 0.1 - branch only when less than 8 grow only when less than 15 neighbouring tips in radius = 20 - - - branching probability = 40 growth speed = 1 |
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Fig. 7. Making the growth of hyphal tips depend on the number of hyphal tips in the neighbourhood. Branching rule the same as in Fig. 6. |
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Interestingly, a similar effect to decreasing the threshold for preventing branching and growth can be achieved also by increasing the radius that defines ‘neighbouring tips’. In this scenario, a sparser structure is produced with longer segments of hyphae before branching is initiated (Fig. 8).
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100 time units: suppose negative autotropism 0.1 - branch only when less than 8 grow only when less than 15 neighbouring tips in radius = 40 - - - branching probability = 40 growth speed = 1 |
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Fig. 8. Increasing the radius that defines the neighbourhood. Growth and branching rules otherwise the same as in Fig. 7. |
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By switching between parameter sets like those described above, it is possible to produce more complex structures. In this example (Fig. 9), three stages were employed:
| first, a parameter set was chosen that produces a dense mycelium, and operated for 100 time units; |
| second, the growth threshold and the size of the radius defining the neighbouring tips were adjusted so that only a few tips continued growth. Furthermore, the probability of branching was reduced to a value close to zero. These settings were operated for 200 time units. |
| Finally, a parameter set that produces a dense globular outgrowth of tips was implemented (for 50 time units). Note that the definition of neighbouring tips is kept large in this parameter set so that tips in the original ‘parent’ mycelium do not resume growth. |
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Parameter set 1 100 time units: suppose negative autotropism 0.1 - branch only when less than 8 grow only when less than 15 neighbouring tips in radius = 20 - - - branching probability = 40 growth speed = 1
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Parameter set 2 200 time units: suppose negative autotropism 0.1 - - grow only when less than 150 neighbouring tips in radius = 75 - - - branching probability = 0.1 growth speed = 1
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Parameter set 3 50 time units: suppose negative autotropism 0.1 - branch only when less than 100 grow only when less than 150 neighbouring tips in radius = 100 - - - branching probability = 80 growth speed = 1 |
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Fig. 9. Compound structure produced by pausing the simulation, changing the parameter settings, and then resuming “growth”. |
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Most parameter settings generate spherical colonies, and that includes a setting in which all controls are removed and growth and branching depend on randomised decisions. In other words, the basic spherical (circular in projection) morphology of the fungal colony arises without the need for global control of that morphology. However, the program does not limit us to spherical end-points.
A thin filament (Fig. 10) can be formed by setting the parameters that prevent growth and branching to high thresholds (i.e. the growth and branching of tips is made highly probable), but then limiting the time for which the tips can grow and branch. (There are two ways to limit the length of branches- first, by limiting the time they grow for, and second, by limiting the length the segments grow to. Both have the same effect.)
To create a number of tips from where the filament can extend any parameter set can be used that generates reasonably large numbers of tips per time unit.
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10 time units: any parameter set that generates reasonably large numbers of tips per time unit.
250 time units: suppose negative autotropism 0.1 - branch only when less than 10 grow only when less than 20 neighbouring tips in radius = 20 stop growing after tip = 10 stop branching after tip = 10 - branching probability = 80 growth speed = 1 |
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Fig. 10. A parameter set that generates a sparsely-branched filament. |
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A more prolifically-branched filament can be produced (Fig. 11) by increasing the threshold for preventing branching and growth (i.e. increasing the density of branches) and by allowing the tips to branch and grow for longer. The branching probability can also be increased to accentuate the effect.
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30 time units: any parameter set that generates reasonably large numbers of tips per time unit.
300 time units: suppose negative autotropism 0.1 - branch only when less than 80 grow only when less than 120 neighbouring tips in radius 80 stop growing after tip = 30 stop branching after tip = 30 - branching probability = 80 growth speed = 1
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Fig. 11. A parameter set that generates a densely-branched filament. |
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Increasing the time that the initial parameter set is run at the start, and then switching to a thin filament parameter set can produce a globular structure with thin ‘exploratory’ filaments emanating from it (Fig. 12).
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100 time units: any parameter set that generates reasonably large numbers of tips per time unit.
200 time units: suppose negative autotropism 0.1 - branch only when less than 8 grow only when less than 15 neighbouring tips = 20 stop growing after tip = 10 stop branching after = 10 - branching probability = 80 growth speed = 1 |
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Fig. 12. Creating exploratory filaments emerging from a spherical mycelium. |
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Implementing the density field hypothesis for branching regulation represents a slightly different way of calculating the density of tips in the locality. The main practical difference between this form of computation and that in the examples discussed above is that when the density field hypothesis is used, local differentiation becomes difficult as the regulation applies to the whole mycelium. For instance, the structure illustrated in Fig. 9 would be impossible using this method, as it cannot prevent growth and branching in the ‘parent’ mycelium while allowing it in the ‘daughters’. Thus, structures are produced that are uniformly distributed with branches and most branching appears to be dichotomous. Fig. 13 shows three examples in which the density field threshold for branching was varied to produce mycelia with different morphologies.
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100 time units: suppose negative autotropism 0.1 suppose density field hypothesis branch if field is less than 0.1 - - - - - branching probability = 40 growth speed = 1
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100 time units: suppose negative autotropism 0.1 suppose density field hypothesis branch if field is less than 0.05 - - - - - branching probability = 40 growth speed = 1
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100 time units: suppose negative autotropism 0.1 suppose density field hypothesis branch if field is less than 0.01 - - - - - branching probability = 40 growth speed = 1
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Fig. 13. Three examples showing effect of variation in the hyphal density field. |
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If negative autotropism is set to zero at the beginning of the simulation (switching this parameter to zero during a simulation produces different results) an interesting growth process can be seen. This can be done with both methods of branching regulation, but the density field hypothesis is supposed in the examples shown here. The interesting feature of the growth of these mycelia is that they grow in one plane at a time. That is to say, a straight, single hypha grows first with many dormant tips along it; then branches grow perpendicularly to produce a 2-dimensional disc-type structure; and finally more branches grow out perpendicularly from that to make a 3-dimensional structure. If a relatively high density field threshold is used, an ellipsoidal morphology is produced (Fig 14).
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100 time units:2 suppose negative autotropism 0 suppose density field hypothesis branch if field is less than 0.1 - - - - - branching probability = 80 growth speed = 1
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Fig. 14. Ellipsoidal morphology. |
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If the density field threshold is reduced somewhat, the result is a rod-like structure.
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1000 time units: suppose negative autotropism 0 suppose density field hypothesis branch if field is less than 0.005 - - - - - branching probability = 80 growth speed = 1
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Fig. 15. Rod-like morphology. |
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Again, switching between parameter sets in the course of a simulation generates interesting compound morphologies. For Fig. 16, three parameter sets were used:
| First, negative autotropism was set to zero and the density field hypothesis supposed, with the threshold for branching set reasonably high (i.e. branching is likely). This produces a short, straight hypha with many (dormant) tips in 90 time units. |
| Second, once the tips on the hypha begin to extend perpendicularly, the branching threshold and probability are lowered so that long branches are grown for 110 time units. |
| Finally, the branching threshold and probability are raised to very high values and the tip growth is limited to 25 time units. This causes dense branching around the growing tips. |
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Parameter set 1 90 time units: suppose negative autotropism 0 suppose density field branch if field is less than 0.1 - - - - - branching probability = 80 growth speed = 1 |
Parameter set 2 110 time units: suppose negative autotropism 0 suppose density field branch if field is less than 0.005 - - - stop branching after tip = 50 - branching probability = 20 growth speed = 1 |
Parameter set 3 25 time units: suppose negative autotropism 0 suppose density field branch if field is less than 0.5 - - stop growing after tip = 25 - - branching probability = 80 growth speed = 1 |
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Fig. 16. Compound morphology produced by three successive parameter sets using regulation by the hyphal density field. |
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For the structure shown in Fig. 17, an ellipsoid mycelium, as described in Fig. 14, was grown initially for 200 time units. Then the parameters were switched to a set that produces densely branched filaments (as in Fig. 11) for a further 100 time units.
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200 time units: suppose negative autotropism 0 suppose density field branch if field is less than 0.1 - - - - - branching probability = 80 growth speed = 1
100 time units: suppose negative autotropism 0.1 - branch only when less than 50 grow only when less than 80 neighbouring tips in radius = 50 stop growing after tip = 20 stop branching after tip = 20 - branching probability = 80 growth speed = 1
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Fig. 17. Compound morphology produced by two successive parameter sets. |
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Now, why don’t you try the program for yourself? If you copy the experiments described above you will get similar results (but remember, each experiment “grows” a new and unique “colony”, so you’ll never get exactly the same).
But you could think up your own experiments, couldn’t you?
Close the window to return to the computer model