Results
We simulated the success of establishment of the pathogen population
during colonization of a new host and, when pertinent, the evolution of
its compound phenotype under the new selective pressure. Parsing these
components allowed a better understanding of the individual and combined
influence of distinct rates of novelty emergence (μ ), propagule
size (N0 ), and reproductive rate
(b )_ on the success of establishment of a new association
and the behavior of the fitness and size of the population of pathogen
following colonization of a new host resource. All analyzed parameters
were evaluated under variable distance of the propagule compound
phenotype from the host (d0 ) and revealed their
influence on the probability of establishment of the pathogen population
in the new host (Fig. 2). For a single propagule
(N0 =1), the increase of d0gradually reduces the probability of establishment (Fig. 2a and b) –
which was an expected result since the survival probability decays
following this distance (black curves in Figs. 2 represent Eq. 1 for the
propagule, di,n =d0 ). For
the pathogen population to colonize the new host, it needs to survive
successive selection events - therefore the probability of establishment
is lower than the survival probability recovered for a single colonizing
individual to persist until the first reproduction (the black curve in
Figure 1).
Greater reproduction rates (b ) favor the pathogen establishment
(Fig. 2a). As b increases, the establishment success approaches
the probability of one individual surviving the selective forces of the
new host species (in Fig. 2a; compare non-black probability curves
approaching the black curve as b increases). For high brates (e.g. b = 7.5), the probability of establishment of the
pathogen population will be the same as that expected for a single
individual surviving until the first reproductive event of the
simulation - and the probability of survival will depend only on the
effect of d0 .
Only high novelty rate values (10-2 and
10-1) had a measurable effect on the populational
probability of establishment - all other variations of novelty rate had
practically the same low effect on the probability (Fig 2b). For novelty
rates between 0.0 and 10-3, the probability of success
practically did not differ, reaching 0 for d0 \(\approx\)1 (propagule compound phenotype app. one standard deviation
distant from the optimum imposed). The effect of the increasing novelty
rate between these values is more evident on the population growth; the
population reaches the carrying capacity about twice faster whenμ =10-4 than when μ =0 (Fig 3). Less than
10% of establishment success was detected for non-synergic simulations
when d0 = 2, despite the novelty rate (Fig. 2b).
Simulations have shown that a small increase in the propagule size (from
1 to 10) greatly expanded the diversity of compound phenotypes which
resulted in a probability of success greater than 90% for pathogens
with a d0 <0.9 (Fig. 2c). For larger
propagule sizes, this success extends up tod0 \(\approx\)1.2. The propagule-size effect was
significant for the survival rate of the compound phenotypes that do not
meet the optimum imposed by the host, maximizing the survival rate of
the neighboring phenotypes as the propagule size increases. This high
probability effect quickly diminishes, depicting a cliff-like pattern
for phenotypes survival probabilities higher thand0 \(\approx\)1.2, independent of the propagule
size.
Finally, the simultaneous maximization of all parameters (B=7.5;μ =0.1; N0 =200) resulted in a synergetic
effect on the probability of success of colonization (Fig. 2). Under
this scenario, even host lineages representing distant resources
(resources that are less compatible with the pathogen
requirements/capacity) have a high probability of colonization, far
exceeding the probability observed for the populational parameters of
the pathogens tested independently (Fig. 2).
As expected, based in every simulated scenario with a non-null μ ,
the emergence of phenotypic novelties in the generations following
colonization allowed the compound phenotypes to evolve towards and
stabilize around the optimal fitness value imposed by the host (Fig. 3).
The greater the novelty rate (μ ), the faster the evolution
towards the optimum, also increasing the diversity of compound
phenotypes (Fig. 3, μ =10-2). During
simulations, population size rapidly reaches the established carrying
capacity. Even though higher values of μ favors population
growth, the carrying capacity is achieved much earlier than phenotype
stabilization for all scenarios (Fig. 3). Surprisingly, even in the
absence of novelties (Fig. 3, μ=0 ) many simulated pathogen
populations persisted and achieved the carrying capacity in the newly
colonized host species.
Varying rates of the emergence of evolutionary novelties revealed also
an unexpected outcome on the qualitative profile of the populations,
following colonization. High rates of emergence resulted in the
retention of compound phenotypes (variants) present in the initial and
previous populations during populational growth, with correspondingly
larger load (something analogous to the concept of genetic load; Wallace
1970) (Fig. 4a). Lower rates of novelty emergence resulted in
populations that depict smaller phenotypic variability, with greater
loss of pre-existing phenotypes (Fig. 4b). By maximizing every other
parameter, the expansion of phenotypes is even larger, indicating that
the increased maintenance of ancestral phenotypes is also influenced by
other populational parameters besides μ .