# Definition of the effector landscape across 13 phytoplasma proteomes with LEAPH and EffectorComb

## TL;DR

This paper introduces LEAPH, a machine learning tool for identifying effector proteins in phytoplasma bacteria, which cause plant diseases.

## Contribution

The novel contribution is the development of LEAPH, a high-accuracy predictor for phytoplasma effectors, and the EffectorComb app for candidate selection.

## Key findings

- LEAPH achieved 97.49% accuracy in predicting effector proteins in phytoplasma.
- Application of LEAPH identified 2089 putative pathogenicity proteins across 13 phytoplasma proteomes.
- Three effector secretion classes were identified, with LEAPH correctly identifying 15 out of 17 known effectors.

## Abstract

‘Candidatus Phytoplasma’ genus, a group of fastidious phloem-restricted bacteria, can infect a wide variety of both ornamental and agro-economically important plants. Phytoplasmas secrete effector proteins responsible for the symptoms associated with the disease. Identifying and characterizing these proteins is of prime importance for expanding our knowledge of the molecular bases of the disease. We faced the challenge of identifying phytoplasma's effectors by developing LEAPH, a machine learning ensemble predictor composed of four models. LEAPH was trained on 479 proteins from 53 phytoplasma species, described by 30 features. LEAPH achieved 97.49% accuracy, 95.26% precision and 98.37% recall, ensuring a low false-positive rate and outperforming available state-of-the-art methods. The application of LEAPH to 13 phytoplasma proteomes yields a comprehensive landscape of 2089 putative pathogenicity proteins. We identified three classes according to different secretion models: ‘classical’, ‘classical-like’ and ‘non-classical’. Importantly, LEAPH identified 15 out of 17 known experimentally validated effectors belonging to the three classes. Furthermore, to help the selection of novel candidates for biological validation, we applied the Self-Organizing Maps algorithm and developed a Shiny app called EffectorComb. LEAPH and the EffectorComb app can be used to boost the characterization of putative effectors at both computational and experimental levels, and can be employed in other phytopathological models.