[Motivation] Code understandability plays a crucial role in software development, as developers spend between 58% and 70% of their time reading source code. Improving code understandability can lead to enhanced productivity and save maintenance costs. [Problem] Experimental studies aim to establish what makes code more or less understandable in a controlled setting, but ignore that what makes code easier to understand in the real world also depends on extraneous elements such as project culture and guidelines, and developers’ background. Not accounting for the influence of these factors may lead to results that are sound but have little external validity. [Objective]: This study aims to investigate how developers improve code understandability during software development through code review comments. Its basic assumption is that code reviewers are specialists in code quality within a project. [Method and Results] We manually analyzed 2,401 code review comments from Java open-source projects on GitHub and find that over 42% of all comments focus on improving code understandability, demonstrating the significance of this aspect in code reviews. We further explored a subset of 385 comments related to code understandability and identified eight categories of code understandability concerns, such as incomplete or inadequate code documentation, bad identifier, and unnecessary code. Among the suggestions to improve code understandability, 83.9% were accepted and integrated into the codebase. Among these, only two (less than 1%) end up being reverted later. We also identified types of patches that improve code understandability, ranging from simple changes (e.g., removing unused code) to more context-dependent improvements (e.g., replacing method calling chain by existing API). Finally, we evaluated the ability of four well-known linters to flag the identified code understandability issues. These linters cover less than 30% of these issues, although some of them could be easily added as new rules. [Implications] Our findings motivate and provide practical insight for the construction of tools to make code more understandable, e.g., understandability improvements are rarely reverted and thus can be used as reliable training data for specialized ML-based tools. This is also supported by our dataset, which can be used to train such models. Finally, our findings can also serve as a basis to develop evidence-based code style guides. [Data Availability] Our data is publicly available at https://codeupcrc.github.io.