A bittersweet acceptance: Between a manuscript and grief

I was so excited when I received notification that my first first-author research paper was accepted. My excitement quickly turned into sadness with the realization that my co-PI was not seeing our vision come to fruition.

The first time I met Scott Hawley was in my genetics module. As a fresh PhD student at the Stowers Institute for Medical Research, I was eager to soak up new knowledge. Studiously, I had already gone through the recommended reading material, chapters from one of Scott’s books. Post it notes were poking out of my binder containing those pages, revealing where I had written down thoughts and questions. The idea of Scott could be intimidating: an infamous scientist in the fields of meiosis and fly genetics. Quickly it became clear that there was no reason to be intimidated. He had a passion for teaching and supporting others where he could. Shortly after we wrapped up the genetics module, I ran into Scott in the cafeteria where he asked if I would consider rotating in his lab. I was exhilarated because I already had the same idea.

When I started my first lab rotation a few months later, I was brimming with what we in Germany would call “Vorfreude” (engl.: “pre-joy”). I had previously met with Scott and my supervisor Cathy about my project for the next eight weeks. The project they had in mind for me was focused on the synaptonemal complex, a structure I had never heard of before coming to Stowers. The topic sounded exciting and I quickly became fascinated by it. What I came to understand was that in early meiosis, this protein structure forms between the paired maternal and paternal copy of each chromosome (Fig. 1a). If it does not form properly, crossovers between these two chromosome copies cannot be made properly, causing the faulty segregation of chromosomes (Fig. 1b). The lab of Scott Hawley was one of the many labs trying to understand how the synaptonemal complex was involved in these processes.

The Hawley lab, and one of my co-authors specifically, had established a mutated fruit fly. This was nothing out of the ordinary for fly people, we love that stuff! This interesting mutation removed the gene coding for the synaptonemal complex protein Corolla and inserted in that very location the coding sequence for the same protein but from a closely related species. My goal for the rotation was to perform the first characterizations of the effects this gene replacement had on meiosis. Cathy already had a plan for which experiments I should do and with her support I got to work. I made crosses, scored flies, and performed microscopy. Some of the work was old habit, most of it was new to me. Quickly it became clear what we were dealing with: The synaptonemal complex in these flies was disassembling too early (Fig. 2a) and, unsurprisingly, there was a moderate but significant increase in faulty chromosome segregation (Fig. 2b)¹. Where it got confusing (and intriguing!) was that crossover rates on the X chromosome were almost entirely abolished (Fig. 2c)¹. The near complete lack of X chromosome crossovers should lead to a dramatically higher rate of errors in chromosome segregation. Scott loved it and he offered me to join his lab to figure out how these phenotypes arose.

A bit less than 16 weeks later I had made my decision. While I was not going to join the Hawley lab as he first imagined it, I was going to be co-supervised by Scott Hawley and Matt Gibson to work on a joint project. However, we agreed that I could continue working on the project I had begun during my rotation in the Hawley lab. The experiments should have been straightforward but as science often goes, it took longer than we thought to finish. As a clear next step, I looked at the rates of crossovers on a different chromosome. Fruit flies only have four chromosomes: The sex chromosomes X and Y, the two large autosomes as known as the 2^nd and 3^rd chromosomes, and a very small 4^th  chromosome. We needed to look at one of the large autosomes. It would not have made a huge difference as we knew there was differences in how crossovers on the sex chromosomes and autosomes respond to synaptonemal complex defects but autosomes between one another did not show major differences².  Therefore, we decided to look investigate crossovers on the 2^nd chromosome as a representative autosome. While crossovers were almost entirely absent from the X chromosome, the effect on crossovers on the 2^nd chromosome was nowhere near as dramatic (Fig. 2d)¹. These results solved our confusion as to why the chromosome segregation machinery was not overwhelmed by chromosomes lacking crossovers.

Thinking the work was done, I presented the story at an internal seminar. As it was my first talk in front of a larger audience, it was exhilarating, especially because I received a lot of questions at the end of my talk. One of those questions was fascinating: “Have you tried looking at these flies at a colder temperature? That might change its disassembly dynamics.” Scott loved the idea and it was an easy experiment to do. The standard rearing temperature we maintained the flies at was 25 °C. I moved these flies to 18 °C instead and indeed, the synaptonemal complex stayed fully intact (Fig. 3a)¹! So, I went back to the bench to repeat the chromosome segregation and crossover experiments at this colder temperature. Due to the rescue of the early disassembly of the synaptonemal complex, crossovers on the X chromosome were improved but overall crossovers still did not reach wild type levels and chromosome segregation was still significantly higher than in wild type flies (Fig. 3b-d)¹. This was even more exciting as it meant that this synaptonemal complex protein, Corolla, is not only part of the structure itself but directly involved in crossover formation.

I was finishing up some last experiments before writing up the complete story. During that time, on a Friday morning in late January I arrived at the lab and Cathy quickly found me. By the look of her face, I immediately knew that she was bearing bad news. Scott had died that morning. Of course, in good old Scott fashion, he had emailed people from the lab about work related matters earlier that morning just prior to that. I liked that. I think that’s how he would have wanted to go, thinking of his science until the very end. Him being gone did not make any sense. I couldn’t even imagine how Cathy must have felt. She has worked with Scott for over two decades. Even for me, Scott was more than just my co-PI. He was my biggest supporter, always looking for ways to push me and my career further. We shared ideas for crazy experiments and talked about papers we recently read. I had realized already that I wanted to continue studying the synaptonemal complex beyond my PhD, a field I stumbled into by accident. And while the fact that he to his last day suffered from imposter syndrome did not fill me with confidence that I could beat mine, a part of me felt like I could make it because he said so. But without him? I was not so sure anymore.

I don’t remember much of what I did the following couple of months, but I did finish those last experiments. With the support of Cathy and Stacie, another long-term Hawley lab scientist, I was able to write up the story for the first time. Looking back, I will always remember the embarrassing first draft I sent them. Cutting myself some slack, it was my first time writing a scientific paper. What I came to appreciate is that a first draft is better than nothing because editing is a lot easier than writing about something for the first time. After several drafts went through multiple people (thank you all!), the manuscript was done. The first submission was desk-rejected, which was frustrating, but the second attempt went better. One round of very fair but major revisions later and the paper was accepted.

The acceptance of a scientist’s first first-author publication in a peer-reviewed journal is a milestone achievement. Regardless of what your paper is about, you had to overcome a multitude of hurdles to get to this point. When I saw the notification, I was filled with so much joy. I shared the news with my co-authors, colleagues, and friends. Everyone was excited! Then the sadness hit me. The one person who will not be celebrating this achievement with me was Scott. At that point it had been almost exactly nine months since he passed. I had accepted his death though certain situations were still upsetting. For some reason I hadn’t considered that this particular moment would upset me as well. It was his exuberant joy which I was able to channel and get out of this. I could imagine the email he would send in response to the manuscript acceptance and his palpable jubilation when we would have celebrated in person. I always knew that it was not me who was special but that he made everyone feel special in those moments, something that I can only imagine stems from his own imposter syndrome. There are plenty of people who will celebrate the very big papers and talk admiringly about their favorites. Scott didn’t care where your paper ended up and every trainee who went through his lab was “one of his best”. It is exactly that skill of Scott’s to make everyone around him feel important and special, that has made it possible for me to accept those bittersweet moments. If I can carry only a fraction of that faith forward, into how I move through my own work, how I teach, and how I mentor, then some part of Scott’s spirit will continue living through me and hopefully inspire others.  

References

1. Williams, S., McKown, G., Yu, Z., Gardner, J., Staber, C., Gibson, M.C., Hawley, R.S., The synaptonemal complex component corolla regulates meiotic crossover formation in Drosophila melanogaster. Chromosoma 134, 10 (2025). https://doi.org/10.1007/s00412-025-00839-z

2. K.K. Billmyre, K.K., Cahoon, C.K., Heenan, G.M., Wesley, E.R., Yu, Z., Unruh, J.R., Takeo, S., Hawley, R.S., X chromosome and autosomal recombination are differentially sensitive to disruptions in SC maintenance, Proc. Natl. Acad. Sci. U.S.A. 116 (43) 21641-21650 (2019). https://doi.org/10.1073/pnas.1910840116

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