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Edited by Joan E. Strassmann, Washington University in St. Louis, St. Louis, MO, and approved January 26, 2018 (received for review August 1, 2017)
Article Figures & SI
Figures
- Fig. 1.
Increased mortality of low-level–infected ants after heterologous pathogen challenge. Mortality of L. neglectus ants was, overall, higher after a challenge with Metarhizium (A; black lines) than Beauveria (B; gray lines). For both pathogens, ant mortality after challenge was affected by the infection state of the individuals, which was either no prior infection (solid lines) or a low-level infection of a pathogen homologous (dashed lines) or heterologous to the pathogen they were challenged with (dotted lines). Homologous low-level infections resulted in protective immunization for Metarhizium but not Beauveria. Heterologous low-level infection, in contrast, consistently increased ant mortality in both pathogen challenges. Different letters indicate groups that differ significantly in post hoc comparisons using Benjamini–Hochberg correction for multiple testing at α = 0.05. Sample size n = 919 ants. Supporting data are given in Dataset S2.
- Fig. 2.
Modulation of behavior displayed by low-level–infected ants toward contaminated nestmates. (A) Aggression, (B) allogrooming, and (C) poison spraying were performed equally toward nestmates contaminated with Metarhizium (black) or Beauveria (gray) but depended on the ants’ infection state, which was either no prior infection or a low-level infection of a pathogen homologous or heterologous to the pathogen the nestmate was contaminated with. (A) The aggression level of low-level–infected ants was higher compared with noninfected controls but did not differ when the nestmate was contaminated with the homologous or heterologous pathogen. (B) Grooming was performed significantly longer by low-level–infected ants to a nestmate contaminated with the homologous pathogen, compared with both noninfected control ants and low-level–infected ants grooming nestmates contaminated with heterologous pathogen. (C) Poison spraying was essentially absent in noninfected control ants but increased in low-level–infected ants interacting with a nestmate contaminated with the homologous pathogen and was increased further when the nestmate was contaminated with the heterologous pathogen. Mean ± SEM displayed separately for Metarhizium and Beauveria, with the line indicating a nonsignificant difference between the two pathogens; different letters indicate groups that differ significantly in post hoc comparisons using Benjamini–Hochberg correction for multiple testing at α = 0.05. Sample size n = 720 ants in 144 independent replicates. For the number of ants engaging in the particular behaviors see Fig. S3. Supporting data are given in Dataset S3. For the behavior toward noncontaminated control nestmates see Fig. S4 and Dataset S4.
- Fig. 3.
Reduced heterologous pathogen transfer to and superinfection of low-level–infected ants. (A) Low-level–infected ants acquired viable, infectious pathogen from their contaminated nestmates. Overall, higher pathogen transfer occurred when the nestmate was contaminated with Metarhizium (black) rather than Beauveria (gray). For both pathogens transfer depended on the infection state of the ants. While ants with a homologous low-level infection acquired equal amounts of the pathogen than noninfected controls, ants with heterologous low-level infections acquired less pathogen than both the noninfected control and ants with homologous infections. (B) Superinfection load of both pathogens was reduced in ants displaying risk-averse behavioral modulation compared with control ants. Mean ± SEM displayed; different letters indicate significance groups of all pairwise post hoc comparisons after Benjamini–Hochberg correction at α = 0.05. Sample size: (A) n = 820 ants in 164 independent replicates and (B) n = 255 ants in 51 independent replicates. Supporting data are given in Dataset S5.
Data supplements
Supporting Information
- Download Supporting Information (PDF)
- Download Dataset_S01 (XLSX)
- Download Dataset_S02 (XLSX)
- Download Dataset_S03 (XLSX)
- Download Dataset_S04 (XLSX)
- Download Dataset_S05 (XLSX)
- Download Movie_S01 (MOV) - Behavioral repertoire of low-level–infected ants displayed toward a contaminated nestmate. L. neglectus workers that had previously acquired low-level infections by social contact with an exposed individual were observed for their behavior toward a newly encountered, contaminated nestmate (red color marking), from which they could contract either the homologous or the detrimental, heterologous pathogen. Displayed behaviors: (i) four low-level–infected ants removing infectious particles from the body surface of their contaminated nestmate by grooming, (ii) a low-level–infected ant disinfecting a contaminated nestmate by antimicrobial poison spraying (two examples), and (iii) a low-level–infected ant performing aggressive behavior (dragging) against a contaminated nestmate (two examples). Still image from Movie S1: In the upper right a low-level–infected L. neglectus worker disinfects its pathogen-contaminated nestmate by antimicrobial poison spraying [i.e., by bending its rear end (gaster tip) with the poison gland exit toward the contaminated nestmate].
Ants avoid superinfections by performing risk-adjusted sanitary care
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