Laidlaw Bee Facility

Named after the “father of honey bee genetics” Dr. Harry H. Laidlaw Jr., the Bee Research Facility is a part of a larger, as I like to refer to it, Bee Complex located only a few miles away off of Central UC Davis Campus (West of Route 113 for those who might be familiar with the area). The Bee Complex is composed of the Bee Facility, UC Davis Bee Haven garden, a number of smaller research plots, several ancillary buildings for storage and most recently, a set of mobile trailers housing the Davis USDA Bee Lab. But as you know, it is the people that really make the program, and our program at UC Davis is a home to a number of researchers and extension specialists contributing to bettering bee health. This series aims to showcase all of the great work being done by the UC Davis Bee Program teams. But first, a little bit about our history!

The Bee Facility Apiary

While you have likely heard of the Bee Program at UC Davis, you probably don’t know that its impressive history began long before many of us reading this issue of Bee Culture were even born. It is my pleasure to take you on a brief journey of the establishment of the UC Bee Program, as it has been shared with me by the late Robbin Thorp and Eric Mussen (to read a more detailed history written by Kathy Keatley Garvey, please visit https://ucanr.edu/blogs/blogcore/postdetail.cfm?postnum=39191). Lastly, I will briefly introduce you to the current faculty members and their respective research and education programs, which we will expand on in the upcoming 2024 issues of Bee Culture. Many well-known and well-respected researchers and educators have conducted seminal works while at UC Davis. They helped make the UC Davis Bee Program what it is today and my current colleagues and I are eager to carry that legacy into the future. My hope is that you will enjoy reading this series as much as we are enjoying writing it for you.

The People: Early 1900s to Early 2010s
Let me take you back to the early 1900s (yes, I just said that, and yes it makes me feel old as well!), when George Haymaker Vansell (1892-1954) was a student at UC Davis. His interest in insect science led him to become the first instructor to teach an Entomology and Apiculture course at UC Davis from 1920 to 1931, highlighting the need for formal Entomology education. He was titled an Instructor in Entomology while also holding a position as a USDA employee at the Davis Experiment Station. Vansell was particularly interested in the field of plant-insect interactions, and has published a number of bulletins concerning honey bee forage. As avid beekeepers, most of you have probably heard at some point that honey bee colonies can suffer poisoning when foraging on California Buckeye (Aesculus californica). Vansell’s interest in this phenomenon resulted in several publications in which he discusses the symptoms of buckeye poisoning, and together with his colleagues, offers possible solutions for reducing damage to colonies. His work suggests that adult worker bees were usually not detrimentally affected; therefore, creating small colony units containing only adult workers and one frame of brood can lead to production of buckeye honey while simultaneously preserving some of the colony work force if the honey is extracted in a timely manner. Vansell and Todd also suggest that Italian x Carniolan hybrids faired a bit better during the production of the buckeye honey as compared to Italian x Caucasian hybrid colonies, but neither had successful recovery. Interestingly, throughout these research articles there is regular mention of the bears destroying colonies in the Sierra Nevada foothills, much like the beekeepers today still have to deal with these intruders. Prior to his sudden passing in 1954, Vansell had also completed research on fruit tree and seed crops pollination. A scholarship established in his honor has helped support many bee students in their fervent effort to advance the field of apiculture.

Around the same time period, Frank Edward Todd (1895-1969) served as the USDA apiculture research branch head at the USDA Pacific States Bee Culture Laboratory at UC Davis (1931-1942). He collaborated closely with Vansell on projects dealing with honey bee poisonous plants, and has advanced pollination of many crops including seed alfalfa, cantaloupes and tangerines. Perhaps most notably, he has reported observations of honey bee nectar collection on alfalfa and the, now well known, tripping mechanism of the alfalfa flower during attempted foraging by honey bees. While affiliated with the UC Davis Bee Biology Program, he modified the dead bee trap originally designed by Norman Gary, which is known as the “Todd Dead Bee Trap”, and has been used in research on effects of various chemicals on bee mortality. Another USDA apiculturist worth mentioning was Edward Lloyd Sechrist (1873-1953). While working in the USDA Office of Bee Culture, he collaborated with researchers at UC Davis Bee lab on several projects that have included honey gathering and daily colony weight changes due to nectar collection. His most notable contribution to the field of apiculture is the proposition for United States standards for honey in 1927.

You probably noticed that the first researchers conducting honey bee and pollination research at UC Davis were actually most directly associated with USDA. However, in 1931, UC Davis hired John Edward Eckert (1895-1975) as a Professor of Entomology and Apiculture, who also served as the Department Chairman from 1934 to 1946. Eckert is well known for studying the flight range of honey bees and he reported extensive observations on this topic including the observation that honey bees prefer to stay close to the apiary in search of forage, but will fly up to 8.5 miles to the food source if necessary. Honey bee resource constancy was also noted by him. Eckert (affectionately called Eck by his peers and stakeholders) was well respected among beekeepers as he supported their efforts to protect colonies from pesticides, and has completed research on potentially harmful pesticide effects on colonies. He is also credited with pioneering antibiotic use in honey bee colonies for management of bacterial diseases, and spent time in Australia and Europe researching various ectoparasitic mites on honey bees including Tracheal mites (Acarapis woodi). Very apropos to this article, Eckert spent decades as the editor of the California column in Gleanings in Bee Culture. Among his many extension publications is the first edition of the Beekeeping in California, Circular 100 from 1936, which has been updated over the years and is still used by many.

As mentioned before, the facility that is still being used by the Bee Program faculty, has been named in honor of Harry Hyde Laidlaw, Jr. (1907-2003) who joined UC Davis as a Professor of Apiculture in 1947. Laidlaw’s research studying mutations leading to differences in eye color, pigment-free blind drones, differences in wing length, hairlessness and resulting identification of underlying molecular and biochemical pathways, have earned him an unofficial title of “The Father of Honey Bee Genetics”. Arguably most impactful applied technology development, however, was the development of the first functional instrument for insemination of queen honey bees. This was made possible by Laidlaw’s study of the queen morphology, and subsequent realization that the only way the queen can be successfully instrumentally inseminated is if the valve-fold is held away from the median oviduct opening. His discovery has provided the means for successful bee breeding and has revolutionized the beekeeping industry. Northern California bee breeders still speak very fondly of Laidlaw. Speaking to his aptitude for innovation and leadership was his selection as the first Dean for Research in the College of Agriculture at UC Davis. He published several seminal queen rearing and bee breeding books, including my personal favorite Queen Rearing and Bee Breeding, written in collaboration with Robert Page, another alumni of our Bee Program. Lastly, in addition to the Bee Biology Facility being named after him, the Laidlaw family established an endowment in his name and in support of student research.

Robbin Thorp, Norm Gary, Larry Connor at the Bee Facility in February 2016

Fifteen years later, Norman E. Gary joined UC Davis as a Professor of Apiculture with special interest in studying honey bee foraging behavior and mating behavior of queens and drones. He was the first to identify queen mating pheromones, and to observe and describe aerial mating of queens and drones. During the medfly eradication efforts by California Department of Food and Agriculture, Gary began studying the impact of pesticide applications on honey bee health, which led to his design of the dead bee trap, later modified by Todd. Gary is also well known for his contributions to the film industry as he has been an adviser on sets of movies such as “Fried Green Tomatoes”, “My Girl” and “Candyman”, earning him the nickname “The Bee Wrangler”. He even has his own IMDb page. Gary has been retired since 1994, but he still occasionally visits the Bee Facility and even borrows bee colonies for small behavioral experiments. Much like his contemporaries Robbin Thorp and Eric Mussen were not, Gary is not very good at being “retired”, and has since published another one of my favorite book recommendations Honey Bee Hobbyist: The Care and Keeping of Bees (I have been lucky enough to have him sign my copy!).

Joining forces with Laidlaw and Gary, Ward Stanger (1913-2000), an extension apiculturist quickly became a champion for the beekeeping industry. In the late 1960s and early 1970s he published extension works discussing the beekeeping industry in California, and comparing the bee breeding and queen rearing efforts in Northern California versus southeastern Gulf States. Stanger understood the value of optimal nutrition to bee health and need for pesticide protection, readily urging the U.S. government to allow for forage access and stricter pesticide regulations. He has also published recommendations for supplemental feeding of colonies to increase their productivity, and a manual on how to remove honey bees from structures.

Christine Peng and Elina L. Niño, January 2020

In 1975, Christine Y. S. Peng joined the Entomology Department as the Professor of Apiculture specializing in insect physiology. I am sure that at this point many, if not all of you, are aware that antibiotics for management of honey bee bacterial diseases require a prescription from a veterinarian. But I bet you did not know that Peng was instrumental in selecting tylosin as a possible replacement antibiotic for oxytetracycline hydrochloride (Terramycin®) since Paenibacillus larvae started developing resistance to it. Peng has also made invaluable contributions in elucidating gamete physiology laying groundwork for successful cryopreservation of honey bee genetic material. Her research into honey bee nutritional needs has led to guidelines for seasonal feeding regimes, and her interest in parasitology has led her to explore varroa mite physiology and various management strategies.

I am pretty certain that Robert E. Page Jr. and his seminal works in honey bee genetics don’t need much of an introduction to the readers of Bee Culture. Page joined the Department of Entomology faculty in 1989 where he also served as the Department Chair. There is not enough space here to write about his many research accomplishments so I invite you to read some of the hundreds of scientific articles or the four books that he has published thus far. His published works report on fundamental discoveries in honey bee behavior particularly regulation of foraging behavior, population genetics and the evolution of complex social behavior. Despite all his achievements and accolades he remains a refreshingly approachable colleague. His passion for honey bees particularly shines through in one of his latest projects “The Art of the Bee” YouTube channel (https://www.youtube.com/@artofthebee).

Sue Cobey, March 2018

The Bee Program can’t really be talked about without mentioning the contributions of Susan Cobey who was at UC Davis from 2007 to 2012. Cobey is a giant in the field of honey bee breeding and has worked tirelessly for decades to maintain and improve quality honey bee stock in close collaboration with the Northern California Bee Breeders. As a young eager doctoral student just discovering my interest in honey bee queen mating physiology, I deeply valued the opportunity to take the world-renowned Instrumental Insemination (II) course with Cobey while she was still working at UC Davis. Principles of II and many tips and tricks shared with me by Cobey are something I now share with the students in our own II courses. Her sustained efforts to improve the bee stock in the U.S. have led to the establishment of the New World Carniolan Breeding Stock that can be purchased from Northern California bee breeders.

Back in 2014, I joined the Department of Entomology and Nematology at UC Davis as the Extension Apiculturist, and to my delight I was able to spend a significant amount of time in the company of two great pollinator researchers and educators: Robbin W. Thorp (1933-2019) and Eric C. Mussen (1944-2022).

Thorp (To read more about R. W. Thorp, visit: https://ucanr.edu/blogs/blogcore/postdetail.cfm?postnum=30459) joined UC Davis as a Professor of Apiculture in 1964 and his research interests were in pollination behavior of honey bees particularly in almond production. Later on, he shifted his focus to non-Apis bees with emphasis on bee systematics, bee conservation and pollination of vernal pool plants. Bumblebee conservation efforts have been in large part inspired by Thorp’s research and he is cited as the main catalyst for successful petition for listing rusty patch bumblebee as an endangered species. Even though he retired in 1994, Thorp continued to come to work at the Bee Facility every day and continued to work on several projects. I have to specifically recommend two books he co-authored in his retirement: Bumble Bees of North America: An Identification Guide and California Bees and Blooms, a Guide for Gardeners and Naturalists. I am forever grateful to him for his guidance and advice, and for not minding me asking him a million questions while he was patiently identifying drawers-full of pinned bees for dozens of student and postdoc projects.

Eric Mussen – Photo by Kathy Keatley Garvey

Similarly, I will forever harbor deep gratitude and appreciation for Eric C. Mussen (To read more about E. C. Mussen, visit: https://ucanr.edu/blogs/blogcore/postdetail.cfm?postnum=52399). He joined University of California Cooperative Extension in 1976 and quickly became a go-to person for the beekeeping industry in California. As he spent more time immersed within California beekeeping, many others such as government entities, non-profit organizations, commodity boards became reliant on him for scientific and practical information. In collaboration with other UC Davis bee researchers, he conducted applied studies immediately relevant for the contemporary beekeeping industry. Shortly before I came to the University, Mussen retired in 2014. I will always be grateful to him for introducing me to the California beekeeping community, for offering guidance, and persistent willingness to give advice while making sure I become fully integrated within the California beekeeping industry.

It was truly my great honor and privilege to learn directly from two great bee researchers and educators. There is absolutely no replacement for their innovation and ingenuity in tackling challenges plaguing bee health, and I only hope I can serve California stakeholders as well as they have. They are very missed!

The People: 2010s to Today
Currently, the Bee Program in the Department of Entomology and Nematology has three core faculty members charged with conducting research and formal and informal education on bee biology and health. Neal M. Williams joined the department in 2009 where he continues working on wild bee biology, native bee conservation and pollination biology. He is devoted to developing supplemental forage mixes to enhance nutrition of all bees in agricultural landscapes of California, as well as modeling potential risks and benefits to bees within California lands. Brian R. Johnson joined the department in 2012 with a strong background in bee behavior. At UC Davis, he continues to study the genetic basis of bee behavior, bee defenses, impact of number of stressors on bee health, spread of Apis mellifera scuttelata hybrids within California, and occasionally conducts projects involving other insects. Most recently, he has published a book Honey Bee Biology which is bound to become a staple reading for beekeepers and researchers alike, and his second book should be coming out soon, so keep an eye out for it. I joined the department as an extension apiculturist in 2014, and learned quickly that I still have much more to learn. California beekeeping is not for the faint of heart and I am really grateful for the super supportive California beekeepers whose backing has allowed me to develop my research and extension program to an advanced level. My team and I conduct research that is directly applicable to beekeepers, including varroa mite management, improved nutrition and enhanced crop pollination. Extension activities are done by all members of my team and they range from offering beekeeping courses and giving club presentations through the California Master Beekeeper Program, all the way to offering technical services such as bee testing and colony inspections through newly established UC Davis Bee Health Hub. Several other of our UC Davis colleagues conduct bee research and we often collaborate with Rachel Vannette and Santiago Ramirez, as well as the two new USDA Bee Researchers Arathi Seshadri and Julia Fine.

Thank you for letting me take you on this short, yet (you hopefully agree) impressive journey through the history of UC Davis Bee Program. Make sure you stay tuned for the next articles in this UC Davis series, and with the upcoming start of pollination season it seems only appropriate to continue with an article delving deeper into some of the bee health and crop pollination research being done in the E. L. Niño Bee Lab. “See you” next month!

Mrs. Root in her teens

Mrs. Susan Hall Root
By: Nina Bagley

Susan Hall was born in 1841 in Ely, Cambridgeshire, England. Her parents were Daniel and Mary Hall, both born in England. They had three children: Robert born in 1838, Susan born in 1841 and, after immigrating to America in 1848, Mary born in 1850. During 1815 and 1837, Ely was in general depression, an agricultural community with no work to be found. The townspeople had terrible living conditions and Ely laborers could no longer maintain themselves. The city of Ely’s population was growing more rapidly than it was in the surrounding countryside. Infectious diseases plagued the countryside, then another Cholera outbreak began in England in 1848. There was a heavy death rate, increasing mortality between 1841 and 1848. This might be one of the reasons why Susan’s father decided to embark on a journey to America with his family for better opportunities.

In 1848, Liverpool, England was the most significant immigration port in the world. Traveling from Ely, Cambridgeshire to Liverpool was 250 miles. Once arriving at the port, families waited in lines, sometimes for days just to get on a ship to sail to America. The journey could take forty to ninety days (about three months) with unfavorable winds and harsh weather.

When this occurred, passengers would often run short of food. Bread, biscuits and potatoes were provided by the shipping companies. The food was terrible and, at times, spoiled.

This was not a cruise ship. Passengers had about two square feet of space. It was dirty, with extraordinarily little ventilation, not to mention lice and rodents. It was a long, wearying journey for a young girl of eight, not knowing what the future would bring, leaving her homeland for an unfamiliar land.

Susan’s father decided to farm and raise his family in Medina County, Ohio. In the new homeland, the family prospers and became community members, attending church and farming their land. As a young girl, Susan had no idea that she would grow up to be a driving force in one man’s life or that that man would become a part of history in Medina, Ohio, the place her father would choose to bring his family to have a better life.

As a young girl, Susan had a schoolmate named Sara Root who was very fond of Susan. Sara felt that her friend Susan would make a good wife for her brother. Sara’s brother Amos Ives Root was away for the Winter in Westville, Ohio, on the river, staying with a relative while attending high school. Sara wrote to her brother saying she had found one of the sweetest girls in all the world as a wife for him.

It was a little embarrassing for Amos I. Root when the two first met, knowing that this schoolmate of his sister knew what she had written to him. It was true love at first sight for both.

A.I. Root wrote: “Her honest simplicity and childlike innocence impressed me from the very first; and, as a matter of course, we two proceeded to get acquainted; and I, for my part, I fear, carried out the program so well that the dear sister was a good many times forgotten and ‘left out in the cold’” (Gleanings in Bee Culture, 1923, pg. 58).

Amos would walk miles to Susan’s family’s farm in unbearable weather to spend a few hours in her company. He called on her once in the middle of the week and every Sunday night! Both Amos and Susan were attending school at the time. When visiting in those early days, staying late was much the fashion. Susan would politely tell Amos that her father went to considerable expense so she could attend a particular school for girls. She could not participate in her studies if he stayed so late, and how important it was for her to get a good night’s sleep. Amos was reluctant to go home at the proper time. Finally, Susan said one evening: “Sir, it is time for you to go home.”

Amos was offended and declared that if he went home, he would go and never return. Susan said, “All right. If you refuse to listen to the dictates of good common sense, it will probably be better for both of us that you should go away and never come back” (Gleanings in Bee Culture, Jan. 1923).

Susan was petite with a kind spirit who knew exactly who she was and what she desired. Having their first lover’s quarrel, Amos left with his head up high and a stern look to teach her that he, Amos I. Root, was not to be dictated to in that manner!

It was a dark night, and he was walking quickly. His temper was getting the best of him and flaring up, which it had done most of his life. Amos started reflecting on how he acted and started to slow down a bit. It was the voice of reason or remembrance of his good mother’s teachings. This is what his mother said: “Old Fellow, is it not possible you are taking offense at the wise words of the best friend you have on earth, and that, instead of being offended, you should recognize her as the one whose price is far above rubies?” (Gleanings in Bee Culture, Jan. 1923.)

Amos crossed the bridge, realizing his mistake. He felt a cold chill all over him, and he turned around, walking calmly back to Mr. Hall’s farm as he hurried up to the house he had just left. Above the front door was Susan’s window to her room. He picked up a pebble and gently tossed it up against her window. And the window went flying up just as he expected! He was always quite sure of himself, so this is what Amos I. Root said: “‘Sue, I humbly beg pardon. You were right, and I was wrong. Will you forgive me?’ Susan responded, ‘All right. Good night.’ And down went the window!” (Gleanings, Jan. 1923).

Amos thought she would come downstairs and give him a kiss of reconciliation; Susan planned nothing of the kind! It was a turning point in his life. Amos finally proposed that the two should be engaged. But Susan insisted that they both were too young to be getting married; Amos was 17 and Susan was 15. Kindly, she told Amos that he did not have the vitality for marriage. A.I. Root was sickly and frail as a child and as a man was not strong. She wanted to postpone marraige for a few years because she was not ready for marriage and wanted to complete her schooling.

A.I. Root would go off and find his way in the big wide world for the next couple of years, making a name for himself, giving lectures and expanding his mind to innovative ideas. He never let the idea go, knowing the two would marry one day. After his lecture tours were over. He returned to his father’s small farm in the woods where he lived as a young man.

Susan’s father feared that Amos could not make a living. Amos would prove him wrong. A.I. Root took a course in watch-repairing and, at twenty-one, started a watch-repairing shop under the pretentious name of A.I. Root & Co. He then proceeded to call on his true love, Miss Susan Hall. Susan’s father was humbled, and he approved them of their marriage.

Mr. and Mrs. Root after they were married.

Three noteworthy events took place in 1861. Abraham Lincoln’s inauguration on March 4, 1861. The Civil War started on April 9, 1861. And, A.I. Root and Susan Hall were married on September 29, 1861. Amos was twenty-one and Susan was nineteen years old.

As the sun rose that Autumn morning after Mr. and Mrs. Root married, they started with horses and carriage on a honeymoon trip. The two were by themselves. Amos put out his hand to Sue as he called her, and she looked smilingly up into his face while he spoke, “Sue, the agreement between us two that we have just entered into is the most sacred and solemn step in our lives. Let us fully consider the new relations that rest on the shoulders of both of us, and may God help us to bear with each other and to bear with patience the new responsibilities that are going to rest on us two. May we two, through thick and thin, for better or for worse, cling to each other.” It was a boyish speech, but it was honest.

Mrs. Susan Hall Root would become his support, “wise counselor” and confidante throughout their marriage, including her husband’s business affairs. Mrs. Root’s hard work and excellent management of the home helped A.I. Root to meet his obligations when they were starting out as a young couple. They would build a homestead and live in Medina, Ohio.

A.I. Root would become a very influential man in many ways, especially in the world of bees. By 1885, the Root name was recognized around the world. Modest and simple in taste, Mrs. Root always avoided publicity, preferring the background of a beautiful home life she had with her husband and children.

Mrs. Root would spend the next sixty years being there for her husband while being an attentive wife and mother, giving birth to five children in twenty years: Ernest R. Root, 1862; Maud E. Root, 1865; Constance M. Root, 1872; Carrie B. Root, 1878; and, having her youngest child at forty-one, Huber Hall Root, 1883. The two would cling together for better or worse, just as they promised one another.

In August 1865, a swarm of bees passed over the A.I. Root & Co. One of the employees of Mr. Root asked jokingly what would you give me if I caught the swarm? Mr. Root replied, a dollar securely boxed. The young man brought A.I. Root the bees, securely boxed, and collected his dollar; the rest is history.

The Root Factory.

A.I. Root founded his bee company in 1869 in his hometown of Medina, Ohio to manufacture beehives and beekeeping equipment. The company was shipping four railroad freight cars of beekeeping equipment everyday, things were going well! A.I. Root was working sixteen-hour days, which sometimes made it difficult to be around him. He started the magazine Gleanings in Bee Culture on January 1, 1873. The first edition of A.I. Root’s book ABC and XYZ of Bee Culture was published in 1879.

Mr. Root constantly worked and expected everyone around him to work just as hard! But that was impossible because Mr. Root was continually working and doing the work of five men until he would exhaust himself to the point that he made himself ill.

Mr. Root would say: “Had I gone on as an evil and angry spirit prompted me to do and not turn back to apologize to my dear wife, Sue, there would have been no A.I. Root Co. There would have been no five dear children brought up in the fear of the Lord, and there would, in all probability, have been no A.I. Root now dictating these words” (Gleanings in Bee Culture, Jan.1923).

Mrs. Root’s children, at some point, all worked for the family business. She was a devoted mother and the most meticulous housekeeper; dust and dirt were her enemies!

Mrs. Root’s daughter Candice Root Boyden authored an article in Gleanings about her mother. The title was “Mother”: “Sweet and modest as the violet of her nature England. Mother always kept herself in the background; only her husband and children, no matter how much credit they had accomplished, should go to her” (Gleanings in Bee Culture, Jan. 1, 1922).

Candice remembered how, as a small child, her mother spent most of her time in the kitchen preparing delicious, healthy foods for the family. Mrs. Root would spend many hours standing over a walnut table with drop leaves while she prepared the family meals. At the end of the table was a shallow drawer to put spoons, cutlery, and other kitchen items. But Mrs. Root would use the drawer for more important things. Neatly filled with toys, the drawer’s contents revealed her love and understanding of a child’s nature.

Mr. and Mrs. Root having a picnic.

The toys were not store-bought, as her daughter would say they were “Treasures, queer bits of metal and wood, an old steel puzzle made by Father, rubber balls, balls of string, little wooden boxes and a little shallow bowl carved from black walnut. But unselfishness, Mother’s dominant characteristic, is revealed in the fact that the drawer, within my recollection, never held anything to help Mother in her work and save her steps.” (Gleanings in Bee Culture, Jan. 1, 1922).

At the time, kitchens were compact, with all their cooking utensils and drawers close by to save the women steps in the kitchen. Mrs. Root was okay with the drawer, which was full of toys for the children. And she did not mind walking back and forth to the big pantry each time she needed something out of it. Mrs. Root often had very severe attacks of pleurisy throughout her life, weakening her heart. She was not a robust woman and sometimes did not have the energy to run around after the children, so the drawer kept their little hands busy and close to their mother.

Mrs. Root would fill a bowl with water and place it on the floor so the children could sail their boats. In the Winter, she would warm the water so their little hands would not get cold. She loved children and had a nurturing way with them.

Mrs. Root did not have the opportunity to attend college, but she took immense pleasure in her children and grandchildren attending college.

Mr. and Mrs. Root would not accompany their family to hotel dinners. They would not go to formal dinners or parties in their honor, but they loved simple picnics in the parks with family around them.

In 1901, A.I. Root built a cabin in the northern part of the Michigan woods and went there to live with his wife. In the forty years of married life, they would finally work side by side, enjoying each other’s companionship.

A.I. Root did not like wintry weather; the cold bothered him. So, a few years after they built their Michigan cabin, they would make another cabin in Florida, where they spent their Winters. Mrs. Root did not like calling it a cabin, so she called it their cottage.

The Roots in their 70s.

The fare from Cleveland, Ohio to Bradenton, Florida was $57.15 a tourist for a round-trip ticket for the Winter. The Roots loved going to their Florida cottage, but Mrs. Root was always overwhelmed with grief to leave all her children, grandchildren and great-grandchildren every Winter.

In the early 1900s, they would spend their Summers in Michigan and Winters in Florida.

The Root Men: J.T. Calvert, Huber, Allen and Ernest Root.

Their sons, Ernest and Huber, were involved in the business of the A.I. Root Company along with their daughter Maude’s husband J.T. Calvert who was the bookkeeper. Around 1900, Ernest took over as editor for Gleanings and kept the bee part of the company going while Huber, more of an inventor like his father, experimented with beeswax. Under Huber’s guidance, the Root company started making candles at the request of the Catholic Church. The local priest was looking for better quality beeswax for their candles and a wick that burned longer. A.I. Root’s sons were carrying the torch for their father so he never had to worry about money again and could devote his time to Mrs. Root, the Congregational Church, family, bees and gardening.

Huber started the candles for the Catholic Church.

Mrs. Root, being in her seventies, enjoyed being outdoors, working in her garden in Florida, and enjoyed spending time with her husband. She enjoyed picking vegetables from her garden and sharing them with her family and neighbors.

Her children felt their mother was the most perfect and unselfish of anyone they had ever known. Mrs. Root captured the hearts around her. She had enough love to go around and was happy when she could help those that were needy, lonely, widowed or fatherless.

Her tender heart cared for the neighbor’s chickens, cats and dogs and ensured they were all fed and cared for. And if Mrs. Root felt a horse was being mistreated, she would not stand for it. This most definitely caused her misery.

A.I. Root wrote: “May I be pardoned for saying that the dear little woman has most faithfully kept her part of the pledge year in and year out? Oh! What would I give if I could truthfully say, ‘I have done as well, or even approximately as well?’” (Gleanings in Bee Culture, 1917, pg. 297).

Mrs. Root suddenly passed away on the evening of Tuesday, November 29, 1921 in Bradenton, Florida, where she and her husband had maintained a cottage for several years. The Roots had just returned to Florida a few weeks before her death. She was feeling exceptionally well and was particularly happy to visit her good friend for many years, Mrs. Ed Nettleton of Medina, who also vacationed in Bradenton, Florida, for the Winter.

Although the family knew it was coming for some time, her life was swiftly ended; the family felt it was due to her arteries being weak from pleurisy attacks over the years.

Mr. and Mrs. Root with a grandchild.

Mrs. Root’s aged husband was too feeble and was advised not to make the long trip back home with the body of his long-time companion.

Mrs. Root’s funeral was held at the home of Ernest Root, her son, on Friday, December 2, 1921, at the old homestead. Mrs. Root was eighty years old. Mr. and Mrs. Root had their sixtieth wedding anniversary a few weeks before her death. She left behind her husband, five children, eleven grandchildren and four great-grandchildren. Mrs. Root was a Medina County, Ohio resident for over seventy years and a friend to all. Mrs. Root is buried in the Spring Grove Cemetery in Medina County, Ohio.

A few years later, Mr. Root caught pneumonia on his way home to Medina, Ohio from Bradenton, Florida. Being feeble, weak and bedridden for several days, the doctor was called, but Mr. Root, knowing it was time, looked at his son Ernest one last time, feeling at peace. He was ready to meet his maker and join his true love, Mrs. Susan Hall Root. A.I. Root took a deep breath and passed away on April 29, 1923, with his children by his side. A.I. Root is buried beside his wife in the Spring Grove Cemetery in Medina County, Ohio.

Ten to twenty percent of the people fleeing Europe in the 1800s did not survive. Not only did Mrs. Root survive, but she survived childbirth, raising five children, cooking and cleaning, washing and ironing in the mid-1800s, and tending to a husband who was constantly inventing and taking chances.

Mrs. Root’s children would have children, and their children would have children, and it has continued for five generations.

The A.I. Root Company is still in business today. The magazine is still being published but instead of Gleanings in Bee Culture it is called Bee Culture: The Magazine of American Beekeeping. Today, A.I. Root is the largest supplier of liturgical candles for Catholic churches. A hundred and fifty-four years later, Brad Root continues the family tradition like his father and grandfathers before him. He is the fifth generation of the Root family. So, the next time you light a Root Candle, think of Mrs. Susan Hall Root who was a friend to all.

I agree with A.I. Root that there would be no A.I. Root Co. if not for a tenacious young girl, Susan Hall Root!

“There is a great woman behind every great man.”

Nina Bagley
Ohio Queen Bee
Columbus, Ohio

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Found in Translation

An Interview with Dr. Hongmei Li-Byarlay
Associate Professor and Project Director for Pollinator Health, Central State University, Ohio
By: Jay Evans, USDA Beltsville Bee Lab

Where are you from originally?
*I was born in Tianjin, China, and came to the U.S. to study for my Ph.D. in 2002.

How did you get interested in science?
*When I was a sixth-grader, I talked to my uncle and told him that I want to be a scientist! Maybe because I had read so many books on the weird creatures in the deep ocean and stories of UFOs.

Where did you go to school and what did you study?
*I went to Tianjin Normal University for my Bachelor’s degree in Biology and Education (dual degree). My senior project was on the effects of metal contamination on bacteria in garlic roots. Then, I went to Nan Kai University for my Master’s degree in Zoology. I studied micro-moths in Northern China and discovered four new species.
In 2002, I went to Purdue University in Indiana for my Ph.D. in Entomology and studied genetics and physiology of fruit flies with Dr. Barry Pittendrigh and Larry Murdock. In 2010, I started my postdoc training with Dr. Gene Robinson at the University of Illinois at Urbana-Champaign, studying behavioral genetics of honey bees. In 2013, I studied epigenetics and aging of honey bees with Drs. David Tarpy at NCSU and Olav Rueppell at UNC-Greensboro.

How did you start your career after school?
*In 2017, I got an offer from Central State University as a new Assistant Professor of Entomology. CSU had just gained their new status as a 1890 Land Grant Institution with USDA. I was very excited to start my own lab.

Which hot topics are you studying now?
*I am studying 1) the molecular and physiological mechanisms underlying the social behavior and ageing of honey bees, such as grooming behavior, aggression and foraging behavior, 2) active breeding efforts for selection of mite-resistant bees by selecting mite-biting stocks and 3) landscape ecology of pollinators and flowers.

Where have you traveled in your studies of bees and what was most memorable?
*I have traveled to China, Germany, Canada, Puerto Rico and many different states in the U.S. The most striking memories were observing and doing experiments with Apis cerana in China, and my trip to Puerto Rico to see and feel the gentle AHBs in reality. I really enjoyed interacting with all the hives there.

What are the biggest challenges facing beekeepers moving forward?
*The desire to find new solutions for mite management is so high, and there are many new ideas. I just hope we all think of new solutions by integrating the sustainability of our hives and our environment.

What gives you hope? What are the best recent discoveries in bee science?
*The government, bee scientists, beekeepers and non-profit organizations are all working together to find the best ways to help our bees, which showed the most love and funding support from the community.
Three of the most interesting discoveries from our lab are:
1)A new publication on Single-cell dissection of aggression in honey bee colonies. https://www.nature.com/articles/s41559-023-02090-0. We are all so excited to use a new sequencing technology to help us to understand bees in a deeper way.
2)Our lab’s new pub about RNA methylation and discovery of long non-coding RNAs underlying bee aggression https://bmcgenomics.biomedcentral.com/articles/10.1186/s12864-023-09411-4
3)We showed that the mandibles (mouthparts) are different between high mite-biting honey bee workers and current commercial colonies. I am also working on a new manuscript to show the striking comparison of mouthparts between two different species of Apis, in hopes this sheds light on mite defenses. https://doi.org/10.3389/fevo.2021.638308

Any advice for future scientists?
*Stay curious and ask questions!

What are your hobbies and other interests beyond bees and science?
*I like running, reading with my kids, hiking and camping in national parks, and meditation.

Jerry: Dr. Keith Delaplane, I remember the first time I met you long ago, I was driving with Nick Dadant down to see you. I read about you and had seen your picture. As I was parking the car, I saw you walking across the parking lot. I jumped out and said, “Dr. Delaplane, I presume? Since then, we have worked together many times, experienced many things and it has been good over the years.

K: It has Jerry, and from the years perspective, you and I have seen a lot over the years.

J: For Bee Culture readers, I think about what you have done, how you have conducted yourself and it has been exemplary. We all have to start somewhere, so where did you grow up and how did that land you in entomology and bees?

K: That is a good question and one I think about a lot because I have enjoyed my life and my career. As you go through life and experience things, you learn more and more and are reminded more and more that you didn’t get here on your own merits. You stand on the shoulders of your family, friends, communities and colleagues every step of the way. You become aware and grateful. I am forever grateful for my upbringing in North Central Indiana growing up on a farm. My dad and his dad were row crop farmers – corn, soybeans and hogs. It’s a great way to grow up.

J: So you were outside a lot. Were you naturally interested in insects on a flower or a fly in the hog trough?

K: My ticket into entomology and the whole natural world was through honey bees and beekeeping. So, my dad was a farmer, we weren’t plugged into ecology; the idea of ecology was new in the 1960’s and ‘70’s, wasn’t it? Farming had an uneasy relationship with ecology and the natural world. We thought we were partnering with nature but it was different than ecology with plowing, using synthetic pesticides and incorporating fertilizers and all that stuff that today. We know exacts a toll on the environment. I was not really ecologically minded as a kid growing up. I was agriculturally minded.

J: Its all about production, isn’t it?

K: It is! It’s all about production, profits and maximizing efficiency to extract the most we can out of the acreage that we had. In my career though I have seen that shift, a good and real shift which is still occurring today toward farming with ecological principles.

To answer your original question, I was interested in nature and insects. Honey bees fit right into that ecological paradigm and they are a natural bridge into entomology and biology and that is the path I followed. Beekeeping was in my family, my grandfather had bees. It was not unusual back then; most farming families kept a few bees. My father used to help him, but he never really took to it.

Then, when I was a young teenager, my parents gave me a beginning beekeeping kit. At the time I thought it was pretty random, I mean, beekeeping as a hobby? It was pretty rare and thought of as an oddball hobby.

J: But it wasn’t hogs and cattle, right?

K: No, and in fact it became a source of contention at times when my father and I had different plans on how I should spend my Saturday (chuckles). I wanted to work on my bees while he wanted me to help on the farm. Not to detract from my parents’ support of my beekeeping, they paid for all my beekeeping supplies and let me do it.

J: Did you learn on your own or did you have a mentor? It was easier back then, you could put your bees in a box in the Spring then go fishing the rest of the year.

K: Yes it was so much easier then, you only had to check on them a few times then take the honey off. It was a Golden Age! Well, I read all the books, I read the classics like ABC and XYZ of Bee Culture, Hive and the Honey Bee, First Lessons in Beekeeping by the Dadant Family and all its successive editions. I did have a mentor named Mr. Paul Champ. He had 300 hives and was able to make a living on it. So once in a while, when my family was out and about, we’d bring our empty quart jar and he’d fill it up. I remember he kept this big honey storage tank right in his kitchen next to the refrigerator. So he was a natural choice to answer my questions. He made house calls! He was with me the first time I saw a queen. I remember the thrill of seeing that queen! It was real – not a mystery – and in my box of bees! I will never forget that thrill.

Those memories are ancient to me now but they go back to my childhood. That rich smell, that scent is unmatchable by anything else.

J: That smell of aster and goldenrod in the Fall…

K: Yes, that pungency and the fascination in the apiary. Beekeeping is so remarkable at every level and I was thrilled with that as a young person. I had a grasp on nature that my peers did not, so they tended to gravitate to our farm. Beekeeping was so interesting to them.

J: Were you a member of 4-H or FFA?

K: Strangely, no, my father and his family were rather insular. We were mostly on our own. As I pursued through my life and career and I became involved in agriculture and extension, they didn’t really understand what I was doing but they supported me regardless.

J: What was it that you wanted to learn and discover that took you to Purdue and LSU?

K: It’s a beautiful story, Jerry. I married early; in fact, I was married by my last semester as an undergraduate at Purdue. I didn’t have a firm career path in mind. I majored in Animal Science, which again is an agricultural major. So once again, I didn’t know in which world I belonged. I had some ambitions of Veterinary College, but I didn’t have the grades for it. Then, I took a class from a very kind and insightful professor named Dr. Wallace Denton in my last semester in “Marriage and Family Counseling”. Imagine that, it’s not even an agricultural topic! He asked me what I wanted to do with my life. I didn’t know, so he made an appointment with me to talk about it. When we met, he asked me what my interests were. I looked down and mumbled, “Well I like keeping honey bees…” I was a little embarrassed because it was an unusual interest. “Well then,” he said, “you should enroll in graduate school and earn a degree in Entomology!”

So, I threw my hat in the ring and lucked out with LSU. My wife and I packed up our things and moved to Baton Rouge, LA. I thought about that story a lot through my career.

Adults don’t think about their influence on kids; it’s a fine line between meddling and giving guidance. I learned as a father that a tension runs, wondering if you are pushing too far. When do you push and when do you hold back? In 20 minutes, he altered the course of my life!

J: You have certainly had good guidance in your life.

K: Yes, I am very grateful, in fact I have an addendum to that story. I knew his daughter. She lived and grew up in West Lafayette. Fast forward a decade later and we ran into her here in Athens, GA. She is the life of the party here, she’s a local musician. Her parents had multiple occasions to visit her, so I have had many opportunities to tell him how important he has been in my life and that I was very grateful.

J: At LSU, is that where the light came on?

K: Yes, once I got to LSU, everything clicked for me. It was a “Eureka” moment for me. I loved my experience at LSU and at the USDA Lab. My major professor, Dr. John Harbo is a well-known researcher in honey bee breeding and genetics. I was in school with Tom Rinderer, Bob Denko, Anita Collins and others. It was a golden time in my career.

The Department of Entomology at LSU was strong and I had great professors. That, Jerry, is when I realized that honey bees are more than just agriculture, and biology is more than agriculture and where honey bees fit into just about everything. If I had to do it all over again, I would still stick with Biology.

J: So from there you went to University of GA?

K: Yes, so it was rather word of mouth, but John Harbo called me and said they had an opening. So, I applied for the job and to my ever-living amazement, they interviewed me and offered me the job.

I may have been the last of the generation in which I moved from earning my Ph.D. straight to a tenure track position in a university. Now, students have to get one or a series of jobs as a post doc before they land a faculty position.

I have a multiple appointment in extension and research. I’ve had some teaching appointments off and on but mostly I have been in the field, the laboratory and in direct contact with beekeepers. It was good.

I have written many times that honey bees are a window to the world and it is a good metaphor. They bridge so many domains of human activity. I can’t think of any animal or any field of agriculture or a hobby that bridges so many separate spheres of human activity. Honey bees are remarkable and I am honored and pleased to work with them.

J: When you received your tenure track position, what went through your mind? What was your goal?

K: I realized that I had been treated well and given special privileges and opportunities, so, “Don’t blow it, Keith!” (chuckles). I have never been one to give myself the benefit of the doubt, and have always had self-doubts, so I was motivated by fear. I applied myself and went through the steps I had been taught to do research and “pretended” to be a good researcher. I figured, if I worked on it, I would eventually become a good researcher. You go through mental gymnastics and after a while it sticks. It’s hard for me to be prideful.

J: That is certainly a motivator! How long were you at UGA and how many students did you have?

K: I was at UGA for 33 years and had seven graduate students under my direction and sat on 15 committees. I also had two post docs. I’m grateful to watch a young mind advance from an elementary mind to one who understand and has skills maybe above your own.

J: But that is the goal, isn’t it?

K: It is the goal! An economy of justice occurs but as you become older, you are happier with that arrangement. I’m happy that they get the glory they deserve, you want to give of yourself to give them that recognition. The saying goes, “youth is wasted on the young”, and while I appreciate the humor it is not exactly true. As you learn life’s lessons and you are managing your own interior life, you become better at contextualizing your own ego. Having an ego is important, but your opinion is not the only one and may not be correct. You rarely know the entire story. That is part of the challenge. This is certainly true with managing honey bees. Beekeeping and beekeepers are interesting because they are so diverse in their understanding.

J: I wish someone would invest say, $50,000 to study the profile of beekeepers. That would be very interesting.

K: I am authentically curious about that. Are goat farmers or gardeners like that or are we a unique creature? Beekeepers are an interesting slice of humanity. We are somewhat self-selecting because we like nature but otherwise we are all over the board with social skills, income level, religious and political views, and education. This makes it difficult to navigate a bee meeting. (laughs)

J: Tell me, what has been your biggest success and a failure that you regret?

K: Well, I was inspired the first few months on the job which is probably typical, but I happened upon the good fortune of serendipity to have a desire to record inspections in the yard for extension purposes. The producer was on staff at UGA and an affiliate with the local public broadcasting station. He saw the value in my recordings and published it for a three year TV show. That really put me on the map and enabled me to travel more for talks and demonstrations. I was invited to speak all over the country, in fact the world. This helped with my early academic promotions.

Next, I teamed up with Dr. Mike Hood at Clemson University. He and I were the closest colleagues with a shared appointment in honey bees in the USA as the two universities are only 60 miles apart. We collaborated a lot. We worked on Varroa mite IPM and published the first economic threshold for Varroa mites which was badly needed. We also ran a series of studies using non-chemical techniques to control Varroa. This occupied about 10 years of the middle of my career.

Unfortunately though, it never became overly successful. This falls in the category of a failure. I think we are overly optimistic to think that bees can develop a true resistance to Varroa mites and the pathogens that result from their damage to the bees. We can produce honey bees with some hygienic traits to remove Varroa mites from the colony, but we are using defense mechanisms that were co-opted to defend the bee from something else, for instance, chalkbrood was the first genetic trait found, in which bees would remove infected mummies from the hive, and the detection and removal of pupae infected with AFB. The overwhelming weight of the data have not come up with a ringing solution for Varroa mite control through genetic resistance.

J: When a queen is mating with 14-20 drones, you don’t know the ancestry of the drones or the queen.

K: True! Traditional animal husbandry doesn’t apply with honey bees; it is much more difficult to think in those terms. Honey bees have multiple mating, how it uses genetics to advance the colony is complicated.

That is why I am so interested in polyandry (multiple mating) which is now the focus of my research. It is successful and provides very basic research. It is going to benefit the fundamentals of science. Also, some of my students and I have conducted a great deal of research on blueberry pollination. It is easy to convert that basic science to deliverables that will help growers. You can use it to adjust the number of colonies needed in the orchard, for example.

Science is self-adjusting, some of it will stand the test of time, while some will fade away.

J: Welcome to life, eh?

K: Yes, science is not a mechanism for finding answers to solve life problems reliably, but it is the best mechanism humanity has and is a necessary part of the process. Some studies fail and are never reported, thus repeated over and over.

J: Would you please provide some advice to BC readers?

K: If you dig into honey bee biology and appreciate the organism, you will enjoy it even more. Reading and finding clues to keep honey bees healthy will give you more pleasure, improve colony health and increase your profit line. Honey bees are a bottomless pit of wonder, curiosity and amazement.

I am working on a book which will be out next year, called Honey Bee Social Evolution, published by John Hopkins Press. I wrote about the similarities between different organisms and the honey bee colony as a unit. They all have the same dynamics. Also read Hilda M. Ransome’s book, The Sacred Bee in Ancient Times and Folklore. It was written in the 1930’s but it is a classic. She wrote about how honey bees have been used worldwide in literature, religion and poetry for centuries. There is no end to this insect and what it can give you.

Figure 1. Erin Rupp of Pollinate Minnesota, Anne Turnham (center) and Ana Heck, formerly of University of Minnesota Bee Squad, now Michigan State University Extension educator for Apiculture ready for a day of honey bee outreach.

This year we decided to interview beekeeper Anne Turnham. Even though she doesn’t like to talk about herself and would have rather been on a walk with her best friend and golden retriever Birdie, we figured we’d get her out of her comfort zone because we think that you all will find her story interesting. It’s hard to make a living with bees, but Anne has found really creative ways to center her love for bees in her life and career. Besides keeping her own colonies, she started and runs her own honey label business and works for the UMN Bee Lab’s Bee Squad doing graphics, visual outreach and educational videos.

Q: What first got you hooked on honey bees?
A(nswer/nne): It was when CCD was making the news. I wanted to find out more and just understand what was going on. I took a class, which was mostly inside, but we did get to go outside into an apiary and stand next to the bees. I found it so fascinating that I decided to become a beekeeper by the end of the weekend.

Q: Were you afraid of bees at all when you first started?
A: No. I’d worked with cockroaches in a past job, so bees were less scary. I have a background in Biology and Chemistry and have always loved nature and understanding nature. So, once I decided to get into beekeeping, I devoured every book and class about bees that I could get my hands on. If I was going to do it, I wanted to do right by my bees. Then, in return, my honey bees became this way for me to connect with nature while being at home with my three kids.

Q: What have you learned from the bees over your years of beekeeping?
A: How connected we are to the health of habitat. Bees are such a direct window into what is happening in the environment. They tell you if the environment is healthy. With hive loss, I could directly trace that onto the landscape. Through bees, I learned how closely we are connected to habitat, and how important it is to plant for bees.

Q: Okay so you run your own honey label business and you do the graphics for all the Bee Lab’s educational materials. How did you go from biology to design?
A: I’m self-taught. Learning graphic design was a means to do what I wanted to do. And what I wanted to do was to help bees and beekeepers. After I fell in love with bees, I fell in love with the beekeeper community. They care about insects, these tiny little insects, which just is a testament to how much they care about the world and all living things.

Q: Tell us a bit more about your honey label business*.
A: I started making honey labels for my own honey that I produced along with two of my friends with whom I shared an apiary. I learned Photoshop. Then I put a little blog out with some designs for my friends, but I did not anticipate that it would turn into a business. But there was a need, and people found out. Honey producers are really proud of their honey and they were looking for labels that reflected their pride and their unique businesses. My first order was actually from California. Someone put my designs on a bigger blog, and then suddenly I was getting a ton of customers!

Q: You work for the UMN Bee Lab doing visual education and science communication. What’s your philosophy or goal with this type of work?
A: I tried to volunteer for the squad because I thought they were the coolest. But instead they hired me as a beekeeper. As time went on I started doing more and more graphics work for the team. My first project was designing the labels and instructions for our Varroa Mite Testing Kit.

As to what my goal is, my goal is to help scientists and pollinator educators put content out in the world that will help bees and in turn help the health of the planet.

Q: You have a special talent for turning complex information into very accessible visual descriptions. What happens in your head when you start thinking about how to communicate a given topic visually?
A: I listen a lot and let things percolate before jumping in. My husband David calls it “cluing it.” I always win at the game Clue because I take copious notes and never miss a detail. It’s not that I’m competitive. It’s more like I want to fix a puzzle. I use my “cluing it” energy to puzzle out how to visualize a concept. I ask a million questions because I want to hear the way [other people] understand things. Another important part of my process is going for long walks. My mind needs to wander.

Q: What are infographics and why are they an important communication tool?
A: Infographics are basically simple illustrations, typically one or two colors paired with text to give you a snapshot of what a longer text is about. They support neurodiverse learners because they cue into what the content is going to be about, giving you a way to digest information or even decide if you want to dig in further. It’s a different way of skimming information you want to learn.

Q: Do you think of yourself as an artist?
A: More like a translator.

Q: Thanks Anne! We’re going to go ahead and call you an artist anyway! We appreciate your sharing the ways in which you support honey bees and the ways in which they support you back!

For more information:
https://beelab.umn.edu/manuals
https://anneturnham.myportfolio.com/work

*A note from the authors: Anne is currently not taking new honey label customers in order to meet the needs of her current clients!

(Below) Figure 2. A graphic imagined and designed by Anne Turnham for the University of Minnesota Beekeeping in Northern Climates manual.

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Found in Translation

An Egg-Level View of Drone Production
By: Jay Evans, USDA Beltsville Bee Lab

Honey bee males, or drones, are belittled but key members of the colony. They also form a test case for one of the most fundamental questions in animals and plants. When there is a distinction between males and females, how does that come about? In bees, as in many other species, development into males or females is not black and white. There are proteins (or in cases like our own species, entire chromosomes) that help set the stage for a cascade of events that determines sex. Most of the time, a single trigger, or ‘sex-determining factor’, starts the male and female cascades, and these cascades generally result in physically different males and females. Both that trigger and the resulting cascade differ across the tree of life, and it is hard to point to common sex-determining factors across the insects, let alone the cascades that generate distinct males and females more generally. Thus, it was a really big deal 20 years ago when a research group in Germany led by Martin Beye won the race to find a plausible sex-determining factor for honey bees (M. Beye, M. Hasselmann, M. K. Fondrk, R. E. Page, S.W. Omholt, 2003. The gene csd is the primary signal for sexual development in the honey bee and encodes an SR-type protein Cell 114, 419–429, https://doi.org/10.1016/S0092-8674(03)00606-8). Just this month, that same group closed the circle by demonstrating the key mechanisms by which this factor kicks off drone versus female production in bees… but first some background.

It is staggering to realize that a European priest, Johann Dzierzon, accurately described the process that leads to male honey bees 180 years ago. He was able to show, experimentally, that queens which had been prevented from mating were exclusively drone layers. Genes were not a thing then, let alone sex-determining genes, but genetics was soon to be a field, and there is evidence that Dzierzon’s insights and experiments helped trigger the appreciation for how genetic variation leads to the diversity we see within species. Dzierzon’s passions included how worker bee body colors reflected both queens and their mates and his careful work likely planted seeds in the mind of fellow priest and apiarist Gregor Mendel, who was starting to conduct the pea breeding experiments that defined his own legacy. A nice recent review by Gene Kritsky builds the case for Mendel’s likely exposure to Dzierzon’s thinking in science circles of the 1850’s and 60’s (Kritsky, G. Bees and Peas: How apiology influenced Gregor Mendel’s research. 2023. American Entomologist, 69, 40-45, doi:10.1093/ae/tmad025). Mendel did not formally acknowledge the assist, and it is unclear whether he would have reached the same conclusions and experiments solo. What is certain is that Dzierzon got pretty much everything correct about honey bee reproduction, marveling at queen nuptial flights and the abilities of queens to take or leave sperm from those flights as they nurtured their developing eggs, “The power of the fertile queen, accordingly, to lay worker or drone eggs at pleasure is rendered very easy of explanation by the fact that the drone eggs require no impregnation, but bring the germ of life with them out of the ovary; whilst otherwise it would be inexplicable and incredible. Thus the queen has it in her power to deposit an egg just as it comes from the ovary, and as the unfecundated mothers lay it; or by the action of the seminal receptacle, past which it must glide, to invest it with a higher degree, a higher potency, of fertility and awaken in it the germ of a more perfect being, namely a queen or a worker bee.”

So, how does recent research close the deal for honey bee sex determination? It was evident that the complementary sex determination (csd) gene identified by Beye and colleagues had a highly variable stretch that shows maybe 20 sequence variants in a given population and 100 overall in the species. If diploid female bees are many hundred-fold more frequent than diploid males (which are generally removed by their sisters during development), a gene with this amount of variation fits the bill as the trigger for sex, but how does it all work? Marianne Otte and colleagues from the Beye lab used several genetic tricks to show that a mismatch for this one gene between two chromosomes is both necessary and sufficient to generate female bees. They used ‘CRISPR’ gene editing of fertilized eggs to nullify sections of that variable region. When this happened, bees that would have developed into females were male. They also inserted a polymorphism into drone-layer queens and those queens then produced viable females. Basically, matches for a tiny region of this one protein were sufficient to bind the protein in ways that changed its effects on the next proteins in the cascade and altered the sex of these bees (see graphic). If one of those amino acids was mismatched between the two gene copies, the resulting poor binding led to a female cascade. That’s a simple mechanism for letting a single gene impact sex determination.

While csd appears to be unique to certain insects with haploid males (bees, wasps, ants in particular), it shows a historical similarity to ‘transformer’ proteins, which are known as key actors in insects with diploid males and females and sex chromosomes (i.e., with sex determination that is more like our own). How the leap was made from traditional sex chromosomes to species with haploid males is another mystery. In a practical sense, researchers are rapidly determining variation at csd across populations at all sizes. There is a cost to colonies when queens are mated to males with matching csd alleles. Even though many such ‘diploid males’ are purged early in development the initial effort to raise them, and patchy brood patterns, can both weigh colonies down. Knowing the exact mechanism by which variation works at this locus allows for accurate screens of breeding stock and larger commercial apiaries to see where adding fresh genes might improve productivity. It’s also really neat to think that every cell of a worker bee (or queen) in your colony carries a tiny genetic difference at one of the thousands of her proteins that defines her life.

Queen banking is the storage of queens individually in cages and placed in a colony to be cared for by worker bees. Northern California queen producers bank excess queens as seasonal demand subsides in the Summer to provide an on-demand supply to beekeepers. This study investigated the potential to bank honey bee queens indoors as an effective system during the Summer. This research compared current Summer outdoor queen banking practices in northern California with banking in indoor temperature-controlled storage facilities. Treatments were separated into three groups: indoor queen banks, outdoor queen banks and a set of unbanked control queens. Three different stocking rates were tested (50, 100 and 198 queens per bank). Queen quality parameters and survival data were assessed using laboratory and field assessment methods. There was no significant difference in queen quality parameters apart from the weight of indoor queens banked at the rate of 100, which were significantly lower than the other banking rates. There was a significant difference in the survival of different stocking rates. Queens banked indoors at a rate of 100 were more likely to survive than other stocking rates, both indoor and outdoor. Queens banked outdoors at the rate of 198 were more likely to survive than other outdoor banking rates. Queens stored indoors had a significantly higher survival of 78 ± 1% than queens stored outdoors with a survival of 62 ± 3%. Indoor banking performed better in quality and survival as compared to outdoor queen banking. Therefore, indoor queen banking has the potential to mitigate increased risk to the valuable Fall queen supply caused by rising, hot, Summer temperatures (Onayemi, 2021; Webb et al., 2023).

The mass storage of mated honey bee queens in reservoir colonies over the Winter was investigated under continental climatic conditions. The mated queens were stored in (a) queenright reservoir (QRR) colonies on a frame with partitioned honeycomb, (b) QRR colonies on frame holding wire screen cages, (c) queenless reservoir (QLR) colonies on frame with partitioned honeycomb and (d) QLR colonies on frame holding wire screen cages. In addition to mass storage, the queens were individually wintered in colonies held in Kirchainer mating hives and in five frame nucleus hives with standard combs as the control group. The queen survival in reservoir colonies was observed from October 2000 to March 2001. No queen survived the Winter in QRR colonies, whereas 16.7% of the queens stored in screen cages and 40.5% of the queens on honeycomb in QLR colonies survived for five months. The queen survival in mating hives and in five frame nucleus hives was 80.0% and 83.3%, respectively. Reproductive performances of surviving queens overwintered in reservoir colonies, mating hives and five frame nucleus hives were evaluated by comparing brood areas and adult bee populations produced in test colonies. There were no differences in numbers of frames of bees and in brood production of queens in test colonies. Thus, mass storage of queens over the Winter did not impair their reproductive performance (Gencer, 2003).

Productivity of honey bee queens in Canada, as measured by area of sealed worker brood and net weight of colonies, was generally higher with queens overwintered in two frame nuclei, than with queens overwintered in a group. Poor acceptance and supersedure of group overwintered queens suggest that this method of storage is not yet acceptable for commercial use. Survival of the nucleus queens was low in outdoor two frame units during the Winter but improved with an indoor system. Overwintering queens indoors in two frame nuclei and outdoors in three to five frame nuclei with supplemental feeding of carbohydrate in late Winter should provide a source of queens which could partially fulfill market demands in the Spring (Mitchell et al., 1985).

Queen cages arranged in a holding frame side to side with wire netting opening for nurse bees to feed queens. Photo from http://dx.doi.org/10.1080/00218839.2023.2165747, used with permission.

Spring imports of queen honey bees are essential to replace Winter colony losses in Canada, but contribute to the spread of treatment-resistant strains of pathogens and undesirable genetic traits. A possible alternative to these imports is the mass storage of queens during Winter. By overwintering a strong colony (queen bank) containing large numbers of mated queens isolated in cages, beekeepers could acquire local queens early in the Spring. In this study, the efficacy of overwintering queen banks at two different queen densities (40 and 80) was tested. In the 40-queen banks (40 QB), 74.2% of queens survived the six month overwintering period, while 42.1% of queens survived in the 80-queen banks (80 QB). When compared to queens overwintered free in their colony, queens from bank colonies were smaller and lighter in early Spring but had similar sperm viability and sperm count. Overwintering queens in banks did not have an impact on their acceptance in a nucleus colony but reduced their oviposition in the initial weeks following their introduction. After several days in nucleus colonies, queens from banks had regained a size and weight similar to that of queens overwintered normally, suggesting that they could perform well over a complete beekeeping season (Levesque et al., 2023).

The production of young, mated queens is essential to replace dead queens or to start new colonies after wintering. Mass storage of mated queens during Winter and their use the following Spring is an interesting strategy that could help fulfill this need. In this study, the survival, fertility and fecundity of young, mated queens stored massively in queenless colonies from September to April (eight months) was investigated. The queens were kept in environmentally controlled rooms at temperatures above and below cluster formation. The results show that indoor mass storage of mated queens can be achieved with success when queen banks are stored above cluster temperature. A significantly higher survival of queens was measured when wintering queen banks at 16°C (60.8°F). Surviving queens wintered at different temperatures above or below cluster formation had similar fertility (sperm viability) and fecundity (egg laying and viable worker population). This study shows the potential of indoor overwintering of honey bee queen banks. This technique could be applied on a commercial scale by beekeepers and queen breeders (Rousseau and Giovenazzo, 2021).

The effect of storage cage level (upper or lower) and its position (peripheral or middle positions) on weight, survival rate and egg laying capacity of queens stored in queenright colonies for various storage periods was studied. Storing mated queens in this way had a significant effect on their weight after 75 days of storage. The means of queen weight were 174.9 and 167.4mg for the upper and lower strips, respectively showing the superiority of the upper one. A significant increase in the mean weight of queens stored in the middle position (172.5mg) was noticed comparing to peripheral ones (169.8mg). All the stored queens had significantly greater weight than their original weight before storage during the different periods of experiment. There were significant differences in the survival rate of mated queens stored in different levels, as the mean survival rate of queens stored in the upper strip (69.3%) was higher than the survival rate of mated queens stored in the lower one (60.1%). The queens stored in middle position attained a significantly higher survival rate (70.7%), than those stored in peripheral ones (58.7%). The overall survival rate was negatively influenced with the increase of storage period. In respect of egg laying capacity measured as sealed worker brood area, queens stored for 45 days produced a significantly larger sealed brood area (875.5cm2) than that produced by queens stored for 75 days (843.2cm2) (Al-Fattah et al., 2016).

Mass storage of queens over the Winter was investigated in colony banks, with each queen held in her own cage within a colony. The major treatments included: (I) a single queen wintered in a small nucleus colony (control); and colony banks with 24 or 48 queens, each held individually in (II) screen cages that prevented workers from entering the cage, but allowed access for queen tending, (III) queen-excluder cages (queen-excluder material has openings of about 55mm that prevent the larger queen but not the smaller workers from passing through the material), or (IV) screen cages until January and subsequent transfer to mini-nuclei until late March. Queens held in excluder cages showed poor survival in all three years of testing, and this system was not viable for commercial use; survival for any one year, or any excluder treatment, was never greater than 25%. In contrast, a two year average of 60% queen survival was found for queens that were stored in individual screened wooden cages within queenless colony banks. No differences in survival of banked queens that were moved between colonies monthly and those that remained in the same colony for six months was found. The success of these systems required the (a) preparation of colony banks that contained large numbers of adult workers produced by maintaining colonies with two queens during the previous Summer, (b) removal of laying queen(s) during the storage period, (c) feeding of colonies well and (d) insulation of colonies in groups of four, to preserve heat and reduce worker clustering in the Winter. Surviving queens from Winter storage systems were virtually identical in quality and colony performance to control queens the subsequent season (Wyborn et al., 1993).

This Egyptian study aimed to investigate some factors affecting stored mated queens’ weight and survival rate as well as post storage performance of these queens after 75 days of storage within queenright colonies. Storing queens in numbers of 20, 30 and 40 had no significant effect on their weight. Mean weight of queen stored in excluder cages (EC) was significantly higher than those stored in screen mesh ones (SC). The mean weight of stored queens in the upper strip was higher than the mean of the lower one. Queens stored in peripheral and middle of a holding frame did not differ significantly from each other. Concerning the queens’ survival rate, the mean survival rate of 20 stored mated queens was the superior rank, while the survival rate of 30 and 40 stored mated queens came next with no significant differences between them. Queens stored in SC had more significant survival rate than those stored in EC. The upper strip had a higher survival rate than the lower one. Queens stored in the middle of a holding frame showed significantly higher survival rate than those in the peripheral. Regarding post storage performance, no significant differences were detected between the brood areas produced by queens stored for 45 or 75 days in the three densities. Queens stored for 45 days and those in the upper level had a significantly higher brood production than those stored for 75 days and those stored in the lower level. Queens stored for 45 and 75 days had no significant differences in supersedure percentages either stored in the three densities, in two levels or in the two positions. The second part of the study involved the storing of virgin queens. This work was aimed to investigate the effect of colony and storage cage type on queens’ survival rate, orphan period on attracted workers as well as storage period and colony strength on queens attractiveness and acceptance. Queens stored in Benton cages (BC) had a higher insignificant survival rate than those stored in emerging ones (EMC). Storing queens in queenless colonies resulted in a more significant survival rate than those stored in queenright ones. Increasing the colonies orphan period attracted more significant workers to old queens. This attractiveness increased significantly with the increase of queen age from three to 30 days old. The younger and older virgin queens were significantly more accepted than the intermediate ones. The average number of attracted workers in nuclei was significantly greater than those recorded in strong colonies and so as the acceptance percentages (El-Din, 2016).

The survival of caged newly-emerged virgin queens every day for seven days in an experiment that simultaneously investigated three factors: queen cage type (wooden three-hole or plastic), attendant workers (present or absent) and food type (sugar candy, honey or both) was studied. Ten queens were tested in each of the 12 combinations. Queens were reared using standard beekeeping methods (Doolittle/grafting) and emerged from their cells into vials held in an incubator at 34°C (93.2°F). All 12 combinations gave high survival (90 or 100%) for three days but only one method (wooden cage, with attendants, honey) gave 100% survival to day seven. Factors affecting queen survival were analyzed. Across all combinations, attendant bees significantly increased survival (18% vs. 53%, p<0.001). In addition, there was an interaction between food type and cage type (p<0.001) with the honey and plastic cage combination giving reduced survival. An additional group of queens was reared and held for seven days using the best method, and then directly introduced using smoke into queenless nucleus colonies that had been dequeened five days previously. Acceptance was high (80%, 8/10) showing that this combination is also suitable for preparing queens for introduction into colonies. Having a simple method for keeping newly-emerged virgin queens alive in cages for one week and acceptable for introduction into queenless colonies will be useful in honey bee breeding. In particular, it facilitates the screening of many queens for genetic or phenotypic characteristics when only a small proportion meets the desired criteria. These can then be introduced into queenless hives for natural mating or insemination, both of which take place when queens are one week old (Bigio et al., 2012).

Even though there are some beneficial aspects of banking queens, there can also be some negative effects on the stored queens. Most of the queen banking techniques involve caging queens in various types of cages. Zajdel et al. (2020) reported that queens stored in “queen banks” suffer primarily from leg injuries after they reviewed numerous studies. Queen injuries associated with caging include: 1) changes in the color of the arolia (pad-like lobes projecting between the tarsal claws), 2) missing leg segments or missing whole legs, 3) arolium deformation and partial or complete loss of arolia and claws and 4) frayed wings and loss of antennae or antennal segments. Leg paralysis, probably resulting from stings, has also been reported. These injuries influence the queen’s motor and sensory abilities and disqualify them as high-quality queens. Even a small number of queens stored in one colony are exposed to injuries from worker bees. Injuries to queens were observed regardless of the age of the workers attending to them and the presence of brood in the bee colony.

References
Al-Fattah, M.A.A.W. Abd, H. A. Sharaf El-Din and Y. Y. Ibrahim 2016. Factors affecting the quality of mated honey bee queens stored for different periods in queen-right bank colonies. Effect of cage level and position on holding frame. J. Apic. Res. 55: 284-291.
Bigio, G., C. Grüter and F.L.W. Ratnieks 2012. Comparing alternative methods for holding virgin honey bee queens for one week in mailing cages before mating. PLoS ONE 7(11) e50150. https//doi.org/10.1371/journal pone 0050160
El-Din, H.A.S. 2016. Honey bee Queens Performance In Relation To Their Long Period Storage In Queenright Colonies. PhD Dissertation, Cairo University, 158 pp.
Gencer, H.V. 2003. Overwintering of honey bee queens en mass in reservoir colonies in a temperate climate and its effect on queen performance. J. Apic. Res. 42: 61-64.
Levesque, M., A. Rousseau and P. Giovenazzo 2023. Impacts of indoor mass storage of two densities of honey bee queens (Apis mellifera) during Winter on queen survival, reproductive quality and colony performance. J. Apic. Res. 62: 274-286.
Mitchell, S.R., D. Bates, M.L. Winston and D.M. McCutcheon 1985. Comparison of honey bee queens overwintered individually and in groups. J. Entomol. Soc. Of British Columbia. 82: 35-39.
Onayemi, S.O. 2021. Indoor Queen Banking As An Alternative To Outdoor Banking, M.S. Thesis, Washington State University, 35 pp.
Rousseau, A. and P. Giovenazzo 2021. Successful indoor mass storage of honey bee queens (Apis mellifera) during Winter. Agriculture 11: 402.
Webb, A., S.O. Onayemi, R.L. Olsson, K. Kulhanek, and B.K. Hopkins 2023. Summer indoor queen banking as an alternative to outdoor queen banking practices. J. Apic. Res. 62: 471-477.
Wyborn, M.H., M.L. Winston and P.H. Laflamme 1993. Mass storage of honey bee (Hymenoptera: Apidae) queens during the Winter. Can. Entomol. 125: 113-128.
Zajdel, B., Z. Jasinski, and K. Kucharska 2020. Are drones injured during storage in own and stranger queenright colonies (Apis mellifera carnica)? J. Agr. Sci. Tech. 22: 453-463.

Clarence Collison is an Emeritus Professor of Entomology and Department Head Emeritus of Entomology and Plant Pathology at Mississippi State University, Mississippi State, MS.

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Dog v. Machine: Identifying the Foul in Foulbrood
By: Jay Evans, USDA Beltsville Bee Lab

American foulbrood has been a consistent, if fortunately rare, curse of beekeepers for centuries. The bacterial agent behind AFB, Paenibacillus larvae, is widespread in managed colonies and yet only rarely triggers symptoms in the form of decayed and highly contagious infected brood. Catching those symptomatic cases early remains a critical goal of bee health management. Many U.S. states benefit from a cadre of bee inspectors who work with beekeepers to identify and act upon AFB infections (e.g., the Apiary Inspectors of America, https://apiaryinspectors.org/). Our own USDA Bee Disease Diagnostics Service, led by Samuel Abban (https://www.ars.usda.gov/northeast-area/beltsville-md-barc/beltsville-agricultural-research-center/bee-research-laboratory/docs/bee-disease-diagnosis-service/), works collaboratively with these inspectors and individual beekeepers to pounce on suspected AFB cases before their damaging shadow increases. While the visual and culturing tools for confirming AFB infections are robust, and that smell is hard to forget, there remains a huge need to rapidly screen apiaries for early signs of infection. The frontiers for this screening are marked by an unlikely pairing of furred partners and incredibly complex machines, and it is worthwhile to see which of these tools will be the most helpful for inspectors and beekeepers.

Starting with the more charismatic tools, trained dogs are sporadically used to help inspectors pin down cases of AFB. The state of Maryland has two such trained dogs, led by Chief Apiary Inspector Cybil Preston (https://www.earthisland.org/journal/index.php/articles/entry/detective-dog-sniffs-out-devastating-honeybee-disease/). These companions certainly have the sensitivity to identify signals given off by diseased brood, but how accurate are dogs with the critical early-stage cases of AFB? A study by Neroli Thomson and colleagues in New Zealand aimed to push the limits of training and detection by dog detectives (Thomson, N.; Taylor, M.; Gifford, P.; Sainsbury, J.; Cross, S. (2023) Recognition of an Odour Pattern from Paenibacillus larvae Spore Samples by Trained Detection Dogs. Animals: 13, 154. https://doi.org/10.3390/ani13010154). Two out of three trained dogs did great, consistently and quickly responding to AFB cues placed in one spot within a twirling carousel of dog dishes (Figure). These dogs were trained using purified spores, so would presumably do great even with empty boxes containing post-AFB scale. Their sensitivity in the indoor arena was at the level of spores found in a fraction of a single infected bee. What needs to be tested is the ability of these dogs to ignore the many other smells coming from a beehive, not to mention the environmental distractions (from stinging bees to nervous beekeepers) they would experience when truly on the job.

Since it is hard to interview a dog to find out the cues they use to detect AFB, I decided to explore the most recent work involving chemical sniffers that separate AFB smells from the large and shifting bouquet that is a beehive. Jessica Bikraun from the University of Western Australia devoted her PhD thesis to this question and already has one peer-reviewed paper showing the power of a machine detective approach (Bikaun, J.M.; Bates, T.; Bollen, M.; Flematti, G.R.; Melonek, J.; Praveen, P.; Grassl, J. (2022) Volatile biomarkers for non-invasive detection of American foulbrood, a threat to honey bee pollination services. Science of The Total Environment, 845, 157123, doi:https://doi.org/10.1016/j.scitotenv.2022.157123). Using readily available Solid phase microextraction (SPME) ‘wands’ as noses, she and colleagues collected air samples wafting from infected larvae in a lab-rearing setup and from larvae embedded in living colonies. Larvae sampled in the lab released 102 identifiable chemicals in the air around them. Of these, 17 were found only in larvae infected with the AFB bacterium, others were common to all bees (they also tested bees with sacbrood and bees that had been killed by freezing, along with healthy controls). How do smells in the pristine lab setting compare to those in actual colonies? The SPME technique, while inexpensive and widely available, is compromised somewhat by the greediness of the SPME noses. If there are overwhelming smells coming from a hive, those molecules might edge out rare diagnostic signals. Field trials identified 116 volatile chemicals from beehives, 17 of which were tied to disease. In the end, only four molecules (2,5-dimethylpyrazine, acetamide, isobutyramide, and methyl 3-methyl-2-oxopentanoate) were indicative of AFB both in lab-cultured bees and in-hive air samples. These four chemicals might form the basis for an accurate and simple test. They are also themselves interesting for possible insights into the disease itself. 2,5-dimethylpyrazine is tagged an agent used by bacteria for inhibiting the growth of other microbes, something P. larvae does exceedingly well. While the research was focused on cues that machines can identify, the team also found candidates for smells that hygienic bees pick up on when scanning for diseased brood. Lactones, for example, are natural compounds found in fruits and elsewhere that are often used as components for food additives. In the airspace of beehives, lactones increased substantially with almost any form of brood stress, from AFB to sacbrood and freeze-killed brood, and the authors suggest these compounds might be another trigger for hygienic responses by nest bees. Sujin Lee and colleagues used a lab-based assay to identify and reconfirm volatile chemicals emitted by larvae suffering from AFB (Lee, S.; Lim, S.; Choi, Y.-S.; Lee, M.-l.; Kwon, H.W. (2022) Volatile disease markers of American foulbrood-infected larvae in Apis mellifera. Journal of Insect Physiology, 122, 104040, doi:https://doi.org/10.1016/j.jinsphys.2020.104040. They then purchased those same chemicals to test for responsiveness by worker bees. Bees reacted to several of the candidates but the authors feel that propionic acid, valeric acid, and 2-nonanone were the cleanest signals of AFB infection. Younger bees reacted more strongly to these smells than did foragers, arguably reflecting the tendency of these younger bees (middle-aged actually) to act as hygienic helpers in the colony.

Both dog noses and artificial noses were shown to be capable of identifying even low levels of AFB in field colonies. The SPME chemical nose seems to have more promise as a consistent service (inspectors could readily collect smells from hives with a SPME wand and then send that wand to an analytical lab) but it would not give the in-the-moment diagnostic provided by dogs and good inspectors. For now, those live inspectors are earning their kibble by advising beekeepers when a problem is likely.

Feeling nostalgic for the days when the word “vaccination” was ubiquitous in casual conversation, we decided to explore honey bee vaccinations by interviewing Annette Kleiser, Ph.D., CEO for the biotech company Dalan Animal Health, which has been granted a conditional license by the USDA for their honey bee vaccine.

If you thought creating a vaccination for a regular body was complicated, try making one for a superorganism! In the case of the vaccine Dalan is developing, every mated adult queen would need to be vaccinated. And right now, they are just working on American Foulbrood (AFB), which, while an outbreak can devastate, remains a less-common concern than other diseases like European Foulbrood (EFB), deformed wing virus, chalkbrood and others. How many beekeepers would think it worth preventively vaccinating against this awful but less common disease, especially if they requeen – and would therefore need to revaccinate – yearly?

Figure 1. The field diagnostic test for AFB is to measure the ropiness of the diseased brood. Photo credit: Heather Chapman

We reached out to Annette to find out more. And she assured us that they were truly in the proof-of-concept phase, with the primary goal of garnering support and interest in the beekeeping world. But we had to wonder, why start with a disease so virulent that challenging bees with it to see if the vaccine worked would be… problematic?

Beekeepers are used to industry pressing technology on them, and most of us have developed a somewhat skeptical stance towards the myriad of supplements, gadgets, and flow hives that are foisted upon us, so we brought that attitude to Annette right away. What was her investment in the beekeeping industry?

First of all, we thought Annette was very cool. She is a biologist by training, then moved to helping researchers to develop products and take their work beyond publications. Annette had done her homework and shared a sentiment that spoke to us: “Honey bees… play a vital role in our food security, mitigating climate concerns and supporting biodiversity. ⅓ of our foods depend on pollination, and honey and other products are critical in our food and pharma industries. Despite this fact, beekeeping has been overlooked too long by traditional animal health to provide modern tools for disease prevention and management. Dalan’s goal is to change this.”

We were on board with their goal of finding solutions to bee diseases that don’t rely on antibiotics or equipment-fed bonfires. As you’ll see in the Q&A, Dalan, like beekeepers, doesn’t believe in silver bullets and instead sees their vaccine as a tool in the larger toolkit of good management, good hygiene, good genetics and good nutrition. After we share her answers to our nosy questions, we will end with some further thoughts and questions that were left pondering…

Q: Talk about honey bee immune response and how it relates to the vaccine.
A: Transgenerational immune priming (TGIP) is a natural process whereby maternal insects protect the next generation of offspring from diseases. Nurse bees will encounter a pathogen in the hive, by let’s say cleaning out larvae that have come down with AFB. The nurse bee will take up some of the bacteria that cause the disease, and some bacteria may end up in the royal jelly that is fed to the queen. When that happens, pieces of the dead bug will end up in the ovaries and stimulate an immune response in the developing larvae making them more resistant to infection once they hatch. Our vaccine builds on this natural process. During the vaccination process, we expose the nurse bees and the queen in a very controlled way to a high dose of dead bacteria to ensure that the correct amount is transported to the eggs.

Q: Currently beekeepers are warned about potential AFB transmission from used equipment or feeding their bees honey from unknown sources. Do you see this vaccine as a game changer for the need of those practices?
A: AFB is so problematic because spores can live in the environment for decades, but other bacteria and diseases can be contracted from the use of honey and other sources. I think caution and good hygienic practices should always be part of the routine to keep colonies safe. However, we see this vaccine as a critical part of an integrated control program for sustainable beekeeping that for too long has relied solely on antibiotics to fend off bacterial diseases.

Q: Hygienic behavior has been helpful to remove AFB infected brood from the colony nest prior to it becoming infectious. Has anybody investigated or discussed the potential efficacy of the vaccine and hygienic behavior in protecting bees from AFB?
A: Honey bees are often called a superorganism where the colony has mechanisms for dealing with diseases and infection on an individual level, and on a colony level. Individually bees neutralize disease-causing agents, through a variety of mechanisms like the cutis, grooming, saliva, antimicrobial peptides and other components of the bee immune system. On the colony level, the disease is eliminated though hygienic behavior, removing sick larvae, food sterilization, propolis, but also increasing the colony temperature and creating a fever or sharing antimicrobial peptides with each other. Our vaccine has to be seen in this context. Vaccination is supercharging TGIP and will be supported by all other superorganism mechanisms of disease prevention. Dalan is currently conducting a multi-year, large scale pilot field trial to study the effect of vaccination in the field, where all is coming together.

Q: Does the current research support the possibility that one vaccine might be able to protect bees from multiple pathogens?
A: Our researchers started looking into this for bacterial brood diseases. We have encouraging early results but that is as much as I can say at this time.

Q: The idea of a bee vaccine is exciting, but beekeepers know to be suspect of things that look like silver bullets. What warnings do you have about the capacities and efficacy of this vaccine?
A: I don’t think there is such a thing as a silver bullet. We still need to provide skilled care for these precious animals. Our vaccine was tested and licensed for one disease. It is the first step towards a new era of disease prevention in honey bees. One day we want these tools to become part of the routine measures to protect honey bees, just like taking our pets to the vet to make sure they are protected and kept safe and ensure that they don’t spread diseases to others.

Q: Are there common misconceptions you would like to address?
A: One of the most common misconceptions is around efficacy. When we develop animal vaccines, and in particular, when working with highly contagious diseases, we had to conduct lab efficacy tests rather than infecting animals in the field. That requires establishing laboratory models that use an extremely high infection pressure forcing a high number of animals to die. This is important to arrive at a statistically significant efficacy result. We infect larvae with 5,000-10,000 spores, while typically less than 200 spores are sufficient to kill larvae in a hive. However, we achieved up to 50% survival in these extreme situations in the lab. If you now think of the field situation where you have a much lower infection pressure in the early stages, plus you add in the superorganism ability for disease mitigation, the assumption is that efficacy of the vaccine to protect from disease will be a lot higher.

Q: The Dalan pipeline has AFB, EFB and chalkbrood. Are the viruses transmitted by V. destructor on your radar?
A: Yes, viruses are very important and tools to protect from them need to be a key part of mite management. Viral vaccines are more difficult to manufacture but now that we know how to develop bee vaccines in general, we are ready to tackle viruses as well.

Q: What else do you want to share with us?
A: We realize that our product is just the start. We must be mindful that modern agriculture must also act responsibly in the use of insecticides and herbicides, and that we are at a tipping point. Our product has to be a part of a concerted/integrated effort to provide bees the best possible environment for them to do their job. This may require not only voluntary reductions in some of the damaging chemical practices, but also potential legislation.

Becky Masterman led the UMN Bee Squad from 2013-2019. Bridget Mendel joined the Bee Squad in 2013 and has led the program since 2020. Photos of Becky (left) and Bridget (right) looking for their respective hives. If you would like to contact the authors with your bee vaccine thoughts, please send an email to mindingyourbeesandcues@gmail.com.

Concluding thoughts:
We felt that Dalan’s desire to gather feedback from experts – beekeepers – was sincere: because vaccinating individual bees seems time consuming, the product would need to be excellent enough to actually save the beekeeper money and labor towards honey bee disease management.

While efficient and effective vaccination still seems a long way off after this interview, we like the mindset that we need to prevent disease as much for our bees as for our neighbor’s bees. If varroa and their virus complex has taught us anything, it is the imperative for this community-minded approach. We like Dalan Animal Health’s investment in the vaccine technology, though beekeeper investment is also needed to move this solution forward. Meanwhile, we can dream of a double disease vaccination that protects against the less common AFB and currently too common and menacing EFB. We would give it a shot!

Resources
Dalan Animal Health https://www.dalan.com/
About American and European Foulbrood https://bee-health.extension.org/american-and-european-foulbrood/

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Mite Drop!
By: Jay Evans, USDA Beltsville Bee Lab

Varroa mites remain the primary source of honey bee colony losses for beekeepers managing from one to 10,000 colonies. Scientists like us and ardent beekeepers are always on the hunt for new ways to reduce varroa damage to bees and their colonies. One intriguing strategy is to make mites simply fall off their adult bee hosts. Short of changing the electric charge of host or parasite, this repellency can come from 1) making hosts less grippy, 2) somehow clogging the incredibly strong tarsi (feet with ‘toes’ and a spongy, oily, arolia) of mites or 3) affecting mite behavior by making them less likely to find safe spots and hang on to their bees for dear life. Dislodged mites are far more vulnerable to hygienic worker bees and might also simply keep falling down to a hostless, hungry and hopefully, short life. This is probably a central reason that female varroa mites spend very little time wandering the combs of beehives unless they are moments away from entering the brood cell of a developing bee. While on adult bees, mites have much incentive to stay right there, whatever their host is doing to drop them.

How do mites adhere to their bees so strongly? When mites are actively feeding on bees they are extremely hard to dislodge, since they are partly under the hardened plates of the bee itself and are gripping with a combination of ‘teeth’ and tarsi. Even while taking a break from feeding, mites know to find safe spots on the bee to attach, favoring locations on the abdomen or thorax that are both hairy and away from swinging legs and biting bee mandibles. How can one make them quit their bees given so many hiding places?

Caroline Vilarem and colleagues in France recently described an ambitious attempt to document the abilities of mites to hang onto surfaces when exposed to organic acids (Vilarem, C.; Piou, V.; Blanchard, S.; Vogelweith, F.; Vétillard, A. Lose Your Grip: Challenging Varroa destructor Host Attachment with Tartaric, Lactic, Formic, and Citric Acids, Appl. Sci. 2023, 13, 9085. https://doi.org/10.3390/app13169085). These scientists deployed one of the coolest low-tech tools to measure how well mites grip onto a surface. While their ‘Rotavar’ sounds both complex and expensive, it is actually a ‘motor-driven rotating toothpick’. Yes, you can do this at home, with a slow (three or so revolutions per minute) motor and a supply of toothpicks. The authors add to that an extremely careful experimental design and complex statistics to show the different abilities of mites to hang onto sticks and bees coated with acetic, citric, lactic, formic and tartaric acids. The results hint at new modes and new candidates for mite control, with the usual caveat that converting a controlled lab assay to field colonies will be challenging.

Schematic diagram of the experimental design and measured parameters. Grip on wood (Rotavar): This method relies on direct contact between Varroa’s arolia and the organic acids. The Rotavar set-up is a motor-driven rotating toothpick used to assess V. destructor’s grip. Grip on bees: the host attachment experiment applies acids to the backs of honey bees to remove mites. T0 represents the administration time for treatments; T + 1 h 30, 24 h, 48 h, or 72 h stand for the time post administration used to make measurements. Figure from https://doi.org/10.3390/app13169085

Some highlights: First, acidity itself does not seem to be the solution. Most notably, even high doses of acetic acid had little impact on the abilities of mites to grab toothpicks and this candidate was quickly discarded. So, what can we glean from the differences between the tested acids? Tartaric acid worked great at dislodging mites from spinning toothpicks but was surprisingly poor at dislodging mites from bees. Prior work suggests that the mode of action for tartaric acid is, at least in part, toxicity towards mites. It is possible that the levels of tartaric acid needed to coat bees with a toxic dose are higher than they are on a relatively smooth and barren toothpick. Toothpicks also attract watery compounds (hydrophilic) while bees are coated with oils and are hence more water-repellent (hydrophobic). Maybe the availability of tartaric acid on toothpicks is higher than it would be on oilier bee bodies. Formic acid also worked much better on the wood surface than on bees, an intriguing insight for a well-used and effective mite control. Formic acid is also known to be directly toxic to mites and their cells, and the authors make clear that both direct toxicity and grippiness are clear and perhaps synergistic targets for mite control. The widely used miticide oxalic acid also wins by being directly toxic to mites at levels that are relatively safe for bees, demonstrating that there are many possible ways to turn organic acids into effective treatments.

Lactic acid came out as the best candidate in the study group for divorcing mites from their bees. This acid worked well at dislodging mites from both toothpicks and bees. Lactic acid does not appear to be highly toxic to mites and instead seems to act by changing the mechanics of hanging on. This is a nice lead for exploring acids with similar qualities for their abilities to both grease the ‘Rotavar’ and make bees a more slippery host. In another intriguing result from this nice study, mites that simply walked across paper holding lactic acid were then less good in future grip tests. What is it about lactic acid that burns, cleans or otherwise insults the complex and surprisingly ‘soft’ tarsi of mites?

If this topic has gripped you, consider reading up on the field thanks to a recent open-access paper on stickiness by graduate student Luc van den Boogaart and colleagues in the Netherlands (van den Boogaart, L.M.; Langowski, J.K.A.; Amador, G.J. Studying Stickiness: Methods, Trade-Offs, and Perspectives in Measuring Reversible Biological Adhesion and Friction. Biomimetics 2022, 7, 134; https://www.mdpi.com/2313-7673/7/3/134). For those of us who have stored ‘Freshman Physics’ in a remote hard drive, they give a clear review of how these forces work across organisms; in their words ‘from ticks to tree frogs’. Maybe their figures and insights will inspire a beekeeper or scientist to dream up a safe, effective route to dislodge mites from bees and prevent them from climbing back on. Pulling in people with a knowledge of physics, or just really good imaginations and the ability to build and deploy Rotavars (imagine how entertaining those can be, a la squirrel spinners… https://www.youtube.com/shorts/nBKb_z4_tGY), can only help in the hunt for new mite controls and healthier bees.

Miss Lillian Love was born into a Quaker family in Marion, Indiana on October 24, 1880. Her family was from Decatur, Indiana. Her father, Granville Love, was born in Indiana. In 1860, he married Nancy J. Gillibrand. Her family came from England and settled in the vicinity of Indianapolis. The two were married on August 11, 1868, in Morgan County, Indiana. Granville was a farmer and ran a huckster wagon, which proved a good business. Mrs. Nancy Love had nine children from 1869 to 1892. She died on January 31, 1936, at eighty-six, in Decatur, Indiana. Lillian’s father, Granville Love, died May 7, 1925, in Guilford, Indiana. All the children would be trained in English and piano at Central Normal College in Danville, Indiana.

Lillian’s parents, Granville and Nancy Love.

Lillian had two years of college, became a teacher, and taught in Indiana and Florida. Women still couldn’t find jobs other than teaching. Lillian and her youngest sister Flossie were involved in women’s rights and equal opportunity for women; they supported women’s rights to vote.

In 1904, Lillian moved to Tacoma, Washington, where she taught for several years. Finding her husband in 1907, she married Jay Levant Hill, who was twenty-five years older than her. Lillian’s husband, Jay, was an inventor who owned his own lumber business. Lillian said that her husband’s “brain was mechanically bent.”

Miss Lillian Love, taken in Washington State prior to her marriage to J. Levant Hill.

Their home would be Mount Shadow Ranch, a two-hundred-acre farm with a charming yellow California-style bungalow, two and a half miles from Elbe, Washington. Surrounding the farm were the bee’s favorite purple hills of fireweed.

Fireweed is a plant that enjoys cool and moist climates and thrives in Pacific Northwest forestlands. It is also considered one of the most prolific plants for honey production, with its nectar having a high sugar concentration. It has a “lightly spicy” or “buttery” flavor.

If you shut your eyes and listen, you can hear the train whistle in the distance as it stops at Park Junction Station in the middle of Mount Shadow Ranch.

1926 – Lillian Hill at Mt. Rainier
Apiary. Image featured in American Bee Journal

“We have some thirty acres under cultivation,” said Mrs. Hill; The rest of the farm is logged-off land which we use to pasture our herd of fifty cattle.”
—American Bee Journal, April 1926.

One day in 1913, an old man came peddling bees. “You have a wonderful place here for bees,” he said, convincing Mrs. Hill to invest in six swarms. Before the older man came along, she had never seen a swarm of bees before. Her six swarms increased and produced so much honey that in 1914 Lillian Hill invested in twenty more swarms, giving her forty hives. That year she harvested 6,500 pounds of pure honey. Lillian had an entrepreneurial spirit and determination to succeed in a male-dominated industry.

Lillian’s marriage to J. Hill.

I don’t know how she accomplished so much. Mrs. Lillian Hill kept a tidy home, raised beef cattle, Duroc-Jersey hogs, geese and ducks, and grew vegetables in the garden. But it would be the bees she loved the most!

Lillian was part owner of the Ranch and owner of the Mount Shadow Apiary. She had a gentle personality. She was independent and had a fire in her eyes that you could see demanded respect.

Being a novice beekeeper found her unprepared. In 1915, she encountered her first obstacle, European Foulbrood that would bring havoc to her beeyard. Words that no beekeeper wanted to hear or experience, the only cure, the dreadful burning of the hives. After that horrible experience with “American Foulbrood,” she only kept a dozen colonies of bees providing honey for her family and neighbors. (American Bee Journal, April 1926.)

“I never camp,” confessed Lillian Hill. “On either the trail of my successes or my failures. I go right on.” That’s her philosophy in a nut-shell.

Although childless, she cared for the children from reform schools, orphan asylums or neighboring farms; she taught the boys and girls everything about beekeeping so they could pay their way through school. She believed that the best and safest way to help any human being is to help him help himself. Particularly, those who needed guidance and education.

In the 1900s, the U.S. was a diverse nation, and its children lived in various circumstances. For years, she had been the leader of the Boys’ and Girls’ Bee Club of Elbe. One of her boys won nearly $80.00 with his exhibits of bees and honey at the Western Washington Fair.

Lillian increased her hives to twenty-six to help one of her boys and it didn’t stop there!

In 1924, to help one of her girls through school, she invested in thirty more hives and loaned them to the girl. The girl lived next door to an abandoned schoolhouse on an acre of ground, which got Mrs. Hill thinking, “I could rent the schoolhouse and land from the school board.”

1921 – Freddie May with his siblings before they were placed in the Washington Children’s Home.

One day in 1924, a young man showed up at the Ranch. His name was Freddie May, and he was born in 1912 in Denver, Colorado. When he was eight years old, he lived in Wenatchee, Washington. His father abandoned the family, and their mother could not care for six children. The children were placed in the Washington Children’s Home in Wenatchee, Washington, in 1921.

Mount Rainier Apiary. Freddie May and Mrs. Lillian Hill.

Freddie somehow got his hands on a newspaper. He came across the ad for a permanent position in beekeeping work. Freddie wanted to learn about the beekeeping business under the leadership of Mrs. Lillian Hill, so he rode on “a bicycle” from Wenatchee, Washington to Elbe, Washington, a hundred and ninety five miles! He was energetic and full of fire and wanted to learn beekeeping.

Lillian took a liking to Freddie and wanted to help him make money to pay his way through school, so she furnished Freddie with plenty of bees on a commission basis of fifty-fifty. In four months, the Colorado cyclist made five hundred dollars for himself.

Freddie would consider Mrs. Lillian Hill his mother and next of kin. Lillian and her husband would become Freddie’s foster parents giving him a home with security. He would attend Eatonville High School and work on the Ranch. He would continue beekeeping and eventually marry and have a family.

During the season of 1925, Mrs. Hill was able to establish the Colorado youth in the schoolhouse helping young boys and girls in need teaching them beekeeping.

In 1926, Lillian Hill had over one hundred and fifty hives of Italian bees, eighty-five at the schoolhouse and sixty-five at home. She produced at least 10,000 sections of comb honey. Mrs. Hill would advertise in the newspapers to get workers “Wanted – an experienced farmer for a permanent position.”

Most of the marketing she did herself in her Buick car. She supplied the best stores in Tacoma and Seattle. “I don’t have to hunt for a market,” declared this energetic woman. In one year, Mrs. Hill raised sixty queens. That was the part of her business that she enjoyed most of all. Mrs. Hill was the president of the Pierce County Beekeepers’ Association for two years.

Mount Rainier

Both triumphs and disasters have knocked often at Lillian Hill’s door on the Mount Shadow Ranch, but neither one ever fazed her. This woman had grit and plenty of it!

Around 1927, Freddie would accidentally run over Lillian’s foot crushing it while she was teaching him how to drive the tractor. An unfortunate outcome was that the doctors had to amputate her leg due to blood poisoning. Lillian had a prosthetic leg from the knee down, but that didn’t stop her. She took it in stride and persevered. In 1929, unfortunately, her husband died. He was the youngest of five and the last of his siblings. He was seventy-one years old. I will say some lives have more trial or tribulations than others, to be sure, but no life is without events that test and challenge us.

In the 1930 census, Lillian is listed as a widow forty-nine years old, with fifty men aged eighteen to sixty-six listed as boarders at the Mount Shadow Ranch and working for Lillian Hill. That’s a lot of men to manage. You would have to have grit and be firm! Among the fifty men working on the farm was Freddie May, the youngest, who was eighteen. His occupation was a Logger.

Not being able to care for the Ranch and losing her husband, not to mention the tractor accident, left her feeling like it would be time to sell the Ranch. Lillian Hill would place the Ranch up for sale.

Advertised in the Tacoma Daily Ledger Sunday, June 23, 1929. “Mountain Shadow Ranch. It is one of the best-stocked Dairy Farms in western Washington, with running water in every field and excellent soil. Forty acres cleared; 120 acres fenced for hogs and cattle; stocked and making money; good seven-room house with school buses to Elbe and Eatonville high school. The farm is a must-see to appreciate it. We will consider small trade—a price of $15,000. Write to Lillian L. Hill for an appointment.”

Family photo of Albert Cook, first wife Nora and their children.

Lillian’s family Bible.

A lot happened in 1930. The Ranch sold, and Lillian Hill married Albert Cook, a widower who worked in the lumber industry. His wife Nora passed away in March of 1929 at the age of fifty-one; they had six children together. Lillian didn’t mind an extended family. She was raising her niece Esther who she adopted at a young age and her foster son Freddie May. After all, Lillian loved children and teaching. Her first husband was in the lumber business so she probably knew Albert Cook.

Albert would marry Lillian in 1930, build apartment buildings and retire from the lumber industry. The two would live in Tacoma, Washington. Lillian’s beekeeping days came to an end, her new occupation would be owner and landlord of her apartment buildings.

Lillian had a very loving relationship for nineteen years with her husband Albert. In May of 1949, Albert passed away at the age of seventy-three. He was buried beside his first wife, Nora, in Tacoma, Pierce, Washington.

The income from the apartments and other investments would give Lillian a comfortable life for the next sixteen years. She lived to be eighty-eight and passed away August 19, 1969 in Tacoma, Pierce, Washington Lillian was a Sixth Avenue Baptist church member. She was buried next to her first husband, Jay Levant Hill.

Freddie May lived to be eighty-three years old. Freddie kept his surname May. He went by Fred (Cook) May, Sr. “Commander” as best by everyone who loved him.

Lillian in her sister Flossie’s backyard. Her dress is purple and black print. She and her sister Flossie always had a matching rhinestone necklace.

Lillian’s youngest sister Flossie, who she remained close with, lived in California. Flossie had a granddaughter Karla who enjoyed her aunt Lillian’s visits. She remembers sitting on her grandmother’s hunter green “davenport” with her aunt Lillian. Her grandma Flossie would sit in her desk chair across the room and the two sisters would talk for hours.

Lillian’s great-niece Karla also remembers how “intriguing” her aunt was. Lillian had blue eyes, was fair-haired and had rosy cheeks. She wore her hair in a braid reaching her waist until one day; she cut it off, curled it up, and put it in a small box for keeping. Karla remembers her Aunt Lillian as sweet but at the same time, tough and gutsy!

Marcus Aurelius was a stoic philosopher. His quote reminded me of Mrs. Lillian Love, her struggles as a woman in the 1900’s and how she put others before her, passing her knowledge about beekeeping on to so many young boys and girls in need. I would like to thank Lillian’s great-niece Karla Babcock for sharing her memories of her Aunt Lillian and grandmother Flossie.

“A life of sacrifice and putting the well being collective first, just like the bees.”
—Marcus Aurelius

Ohioqueenbee
Nina M. Bagley
Columbus, Ohio.

Pheromones are involved in intraspecific chemical communication; however, the glands associated with compounds used in nestmate recognition in honey bees remain elusive. This search is difficult since nestmate cues can arise from both within the colony, and from the environment (Kalmus and Ribbands, 1952). For example, Downs and Ratnieks (1999) found no evidence that honey bee guards used heritable cues; instead, guards appear to rely exclusively on environmental cues to distinguish nestmates from non-nestmates. However, nestmate cues can also be produced by the individual, and thus must be under genetic control (Breed, 1983; Page Jr. et al., 1991). A further factor is that the wax used to build comb in the colony is both produced and manipulated by the bees, which means it may be a medium into which recognition cues are transferred (Breed et al., 1998). Therefore, Breed et al. (1998) stated that no single factor is responsible for nestmate recognition in honey bees; rather, all three factors (genetically determined cuticular signatures, exposure to comb wax, and environmental cues e.g. floral cues) seem to work together (Martin et al., 2018).

Comb wax in honey bee colonies serves as a source and medium for transmission of recognition cues. Worker honey bees learn the identity of their primary nesting material, the wax comb, within an hour of emergence. In an olfactometer, bees discriminate between combs on the basis of odor; they prefer the odors of previously learned combs. Representatives of three of the most common compound classes in bee’s wax were surveyed for effects on nestmate discrimination behavior. Hexadecane, octadecane, tetracosanoic acid and methyl docosanoate make worker honey bees less acceptable to their untreated sisters. Other similar compounds did not have this effect. These findings support the hypothesis that nestmate recognition in honey bees is mediated by many different compounds, including some related to those found in comb wax (Breed and Stiller, 1992).

Breed et al. (1998) investigated how kin recognition cues develop and cue differentiation between honey bee colonies. Exposure to the wax comb in colonies is a critical component of the development of kin recognition cues. In this study, they determined how the cues develop under natural conditions (in swarms), whether the genetic source and age of the wax affect cue ontogeny, and whether exposure to wax, as in normal development, affects preferential feeding among bees within social groups. Cue development in swarms coincided with wax production, rather than with the presence of brood or the emergence of new workers; this finding supported previous observations concerning the importance of wax in cue ontogeny. Effective cue development required a match between the genetic source of the workers attempting to enter the hive, the wax to which they were exposed and the guards at the hive entrance. The wax must also have been exposed to the hive environment for some time. Cues gained from wax did not mask or override cues used in preferential feeding interactions; this finding supports the contention that two recognition systems, one for nestmate recognition and the other for intra-colonial recognition, are present.

Recognition of nestmates from aliens is based on olfactory cues, and many studies have demonstrated that such cues are contained within the lipid layer covering the insect cuticle. These lipids are usually a complex mixture of tens of compounds in which aliphatic hydrocarbons are generally the major components. Dani et al. (2005) tested whether artificial changes in the cuticular profile through supplementation of naturally occurring alkanes and alkenes in honey bees affect the behavior of nestmate guards. Compounds were applied to live foragers in microgram quantities and the bees returned to their hive entrance where the behavior of the guard bees was observed. In this fashion, they compared the effect of single alkenes with that of single alkanes; the effect of mixtures of alkenes versus that of mixtures of alkanes and the whole alkane fraction separated from the cuticular lipids versus the alkene fraction. With only one exception (the comparison between n-C19 and (Z)9-C19), in all the experiments bees treated with alkenes were attacked more intensively than bees treated with alkanes. This led them to conclude that modification of the natural chemical profile with the two different classes of compounds has a different effect on acceptance and suggests that this may correspond to a differential importance in the recognition signature.

Cuticular hydrocarbons (CHCs) function as recognition compounds in honey bees. It is not clearly understood where CHCs are stored in the honey bee. Martin et al. (2018) investigated the hydrocarbons and esters found in five major worker honey bee exocrine glands, at three different developmental stages (newly emerged, nurse and forager) using a high temperature GC analysis. They found the hypopharyngeal gland contained no hydrocarbons nor esters, and the thoracic salivary and mandibular glands only contained trace amounts of n-alkanes. However, the cephalic salivary gland (CSG) contained the greatest number and highest quantity of hydrocarbons relative to the five other glands with many of the hydrocarbons also found in the Dufour’s gland, but at much lower levels. They also discovered a series of oleic acid wax esters that lay beyond the detection of standard GC columns. As a bee’s activities changed, as it aged, the types of compounds detected in the CSG also changed. For example, newly emerged bees have predominately C19-C23n-alkanes, alkenes and methyl-branched compounds, whereas the nurses’ CSG had predominately C31:1 and C33:1 alkene isomers, which are replaced by a series of oleic acid wax esters in foragers. These changes in the CSG were mirrored by corresponding changes in the adults’ CHCs profile. The CSG is a major storage gland of CHCs. As the CSG duct opens into the buccal cavity (mouth), the hydrocarbons can be worked into the comb wax and could help explain the role of comb wax in nestmate recognition experiments.

Worker honey bees are able to discriminate between combs on the basis of genetic similarity to a learned comb. The nestmate recognition cues that they acquire from the comb also have a genetically correlated component. Cues are acquired from comb in very short exposure periods (five minutes or less) and can be transferred among bees that are in physical contact. Gas chromatographic analysis demonstrates that bees with exposure to comb have different chemical surface profiles than bees without such exposure. These results support the hypothesis that comb-derived recognition cues are highly important in honey bee nestmate recognition. These cues are at least in part derived from the wax itself, rather than from floral scents that have been absorbed by the wax (Breed et al., 1995).

Experiments indicated that the most important recognition pheromones are the fatty acids, particularly palmitic acid, palmitoleic acid, oleic acid, linoleic acid, linolenic acid and tetracosanoic acid. These fatty acids are mixed with the wax hydrocarbons from wax glands, molded into comb and then transferred onto the workers as they contact the comb. The result is a colony level signature that varies little among workers in a colony. Newly emerged workers have few external fatty acids or hydrocarbons. Oleic acid is more abundant than the other fatty acids on newly emerged bees, but the amount of oleic acid on the cuticle does not vary significantly among colonies. Newly emerged workers are accepted even though they have no signature yet; the “password” for new bees to be admitted to their colony is apparently the lack of a signal. This conclusion is corroborated by the finding that guards tend to treat sodium hydroxide-washed older bees as if they are newly emerged (Breed, 1998).

The integration of recognition cues is described as follows. Fatty acids and hydrocarbons are components of the wax comb that is produced by the bees. The relative abundances of fatty acids and hydrocarbons in wax varies among colonies, giving them unique chemical signatures. Food odors may also be absorbed by the comb, adding to its uniqueness. Newly emerged bees produce their own hydrocarbon coating, which is modified as they move around the nest by the addition of hydrocarbons and fatty acids from the comb. Of the compounds tested in the laboratory, fatty acids are the most important recognition pheromones, but other, as yet untested compounds may also contribute to the recognition odor. Hydrocarbons have generally been assumed to be the primary recognition pheromones of honey bees. However, none of the major structural hydrocarbons of honey bees (i.e., n-alkanes) yields a positive result in a recognition bioassay, nor do these compounds differ significantly in relative concentration among families of bees (Breed, 1998).

The environmental and genetic components of recognition are difficult to separate even in controlled conditions. Getz and Smith (1983) showed that the honey bee discriminates between full and half-sisters raised in the same hive, on the same brood comb in neighboring cells, thus demonstrating a significant genetic component to the recognition process.

Nestmate recognition information can come from either contact chemoreception or olfaction. Mann and Breed (1997) investigated what role airborne olfactory cues play in nestmate recognition by honey bee colony guards, and how do these signals affect guard orientation and behavior? They demonstrated that airborne cues play a significant role in guard bee recognition of nestmates and non-nestmates. Exposure of a guard bee to the scent of a non-nestmate resulted in increased locomotory rate and changes in the directional orientation of guard bees. Exposure to scent of a non-nestmate did not, however, increase the likelihood that a second non-nestmate would be attacked when placed with the guard. Observations of guard behavior at colony entrances indicate that guards discriminate nestmates from non-nestmates with high efficiency.

Floral oils are an important component of the honey bee’s olfactory environment. Bowden et al. (1998) used laboratory and field tests to determine whether floral oils affect nestmate recognition in honey bees. In the laboratory, newly emerged worker bees, that have not been exposed to comb wax, responded more aggressively to bees that had been exposed to floral oils than unexposed control bees. In the field, guard bees did not respond differently to foragers that had been exposed to floral oils. Floral oils may play a supplementary role in nestmate recognition; however, if they have any effect, it is secondary to cues acquired from comb during development.

Downs et al. (2000) investigated the effect that floral oils (anethole, citronellal, limonene and linalool) have on the probability of nestmates and non-nestmates being accepted by guard bees at nest entrances. Floral oils did not affect the probability of workers, either nestmates or non-nestmates, being accepted by guards. However, the presence of floral oils did increase the time taken for a guard to reject an introduced bee. These data show that guards are sensitive to floral oils but use other recognition cues when assessing colony affiliation.

Honey bees have the ability to distinguish among groups of larvae that are destined to become queens and preferentially rear highly related nestmate larvae over less related larvae that are not nestmates (Page and Erickson, 1984).

Colonies of honey bees from two patrilines (cordovan and dark) were established and observations were made on the behavior shown by the worker bees in rearing queen larvae within their colonies. The relationship among the bees within these colonies was either r = ¾ (super-sisters) or r = ¼ (half sisters). The worker bees showed preferential care to the queen larvae that were of their own patriline. Workers of the cordovan patriline showed a stronger preference for larvae of their own patriline than did the dark workers. Cordovan workers also showed a higher rate of visitation, indicating behavioral differences between the patrilines. These results suggest that kin selection is operating on honey bee behavior used in rearing reproduction (Noonan, 1986).

A honey bee queen is usually attacked if she is placed among the workers of a colony other than her own. This rejection occurs even if environmental sources of odor, such as food, water and genetic origin of the workers, are kept constant in laboratory conditions. The genetic similarity of queens determines how similar their recognition characteristics are; inbred sister queens were accepted in 35% of exchanges, outbred sister queens in 12% and non-sister queens in 0%. Carbon dioxide narcosis (stuper, unconsciousness) results in worker honey bees accepting non-nestmate queens. A learning curve is presented, showing the time after narcosis required by workers to learn to recognize a new queen. In contrast, workers transfer results in only a small percentage of the workers being rejected. The reason for the difference between queens and workers may be because of worker and queen recognition cues having different sources (Breed, 1981).

Boch and Morse (1974, 1979) have shown that honey bee queens can be recognized individually by swarms of bees. They found that marking a queen with shellac-based paint to give her a distinctive odor resulted in workers later exhibiting a preference for any queen marked with that paint. However, their experiments do not show whether the odors used by workers to recognize queens are produced by the queens or are environmentally acquired. In a series of studies concerned with queen introduction into colonies, Szabo (1974, 1977) also found that workers could discriminate among queens, but did not approach the issue of the source of recognition odors directly. It was also found that factors such as the age and weight of an introduced queen could affect worker choice among introduced queens. Yadava and Smith (1971) found that the mandibular gland contents of the queen were important in the release of worker aggression towards an introduced queen (Breed, 1981).

References
Boch, R. and R.A. Morse 1974. Discrimination of familiar and foreign queens by honey bee swarms. Ann. Entomol. Soc. Am. 67: 709-711.
Boch, R. and R.A. Morse 1979. Individual recognition of queens by honey bee swarms. Ann. Entomol. Soc. Am. 72: 51-53.
Bowden, R.M., S. Williamson and M.D. Breed 1998. Floral oils: their effect on nestmate recognition in the honey bee, Apis mellifera. Insectes Soc. 45: 209-214.
Breed, M.D. 1981. Individual recognition and learning of queen odors by worker honey bees. Proc. Nat. Acad. Sci. USA. 78: 2635-2637.
Breed, M.D. 1983. Nestmate recognition in honey bees. Anim. Behav. 31: 86-91.
Breed, M.D. 1998. Recognition pheromones of the honey bee. Bioscience 48: 463-470.
Breed, M.D. and T.M. Stiller 1992. Honey bee, Apis mellifera, nestmate discrimination: hydrocarbon effects and the evolutionary implications of comb choice. Anim. Behav. 43: 875-883.
Breed, M.D., M.F. Garry, A.N. Pearce, B.E. Hibbard, L.B. Biostad and R.E. Page, Jr. 1995. The role of wax comb in honey bee nestmate recognition. Anim. Behav. 50: 489-496.
Breed, M.D., E.A. Leger, A.N. Pearce, and Y.J. Wang 1998. Comb wax effects on the ontogeny of honey bee nestmate recognition. Anim. Behav. 55:13-20.
Dani, F.R., G.R. Jones, S. Corsi, R. Beard, D. Pradella and S. Turillazzi 2005. Nestmate recognition cues in the honey bee: differential importance of cuticular alkanes and alkenes. Chem. Senses 30: 477-489.
Downs, S.G. and F.L.W. Ratnieks 1999. Recognition of conspecifics by honey bee guards (Apis mellifera) uses non-heritable cues applied to the adult stage. Anim. Behav. 58: 643-648.
Downs, S.G., F.L.W. Ratnieks, S.L. Jefferies, and H.E. Rigby 2000. The role of floral oils in the nestmate recognition system of honey bees (Apis mellifera L.). Apidologie 31: 357-365.
Getz, W.M. and K.B. Smith 1983. Genetic kin recognition: honey bees discriminate between full and half sisters. Nature 302: 147-148.
Kalmus, H. and C.R. Ribbands 1952. The origin of the odours by which honey bees distinguish their companions. Proc. R. Soc. Lond. B. 140: 50-59.
Mann, C.A. and M.D. Breed 1997. Olfaction in guard honey bee responses to non-nestmates. Ann. Entomol. Soc. Am. 90: 844-847.
Martin, S.J., M.E. Correia-Oliveira, S. Shemilt, and F.P. Drijfhout 2018. Is the salivary gland associated with the honey bee recognition compounds in worker honey bees (Apis mellifera)? J. Chem. Ecol. 44: 650-657.
Noonan, K.C. 1986. Recognition of queen larvae by worker honey bees (Apis mellifera). Ethology 73: 295-306.
Page, R.E. Jr. and E.H. Erickson Jr. 1984. Selective rearing of queens by worker honey bees: kin or nestmate recognition. Ann. Entomol. Soc. Am. 77: 578-580.
Page, R.E. Jr., R.A. Metcalf, R.I. Metcalf, E.H. Erickson Jr. and R.L. Lampman 1991. Extractable hydrocarbons and kin recognition in honey bee (Apis mellifera L.). J. Chem. Ecol. 17: 745-756.
Szabo, T.I. 1974. Behavioural studies of queen introduction in the honey bee 2. Effect of age and storage conditions of virgin queens on their attractiveness to workers. J. Apic. Res. 13: 127-135.
Szabo, T.I. 1977. Behavioural studies of queen introduction in the honey bee 6. Multiple queen introduction. J. Apic. Res. 16: 65-83.
Yadava, R.R.S. and M.V. Smith 1971. Aggressive behavior of Apis mellifera L. workers towards introduced queens II. Role of mandibular gland contents of the queen in releasing aggressive behavior. Cand. J. Zool. 49: 1179-1183.

Clarence Collison is an Emeritus Professor of Entomology and Department Head Emeritus of Entomology and Plant Pathology at Mississippi State University, Mississippi State, MS.

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Found in Translation

Sweet and Sour Honey
By: Jay Evans, USDA Beltsville Bee Lab

There are many ways that honey bees improve our diets but honey consumption was an early reason to wrangle this species. The taste for honey persists today around the world, sustaining sideliners, families and large corporations in many parts of the world. It is also widely known to soothe and improve relations with neighbors, in-laws and bosses. With any high-value product, there is a risk of inadvertent or purposeful false advertising.

One honey quality trait that is easy to control is water content. Small-scale beekeepers routinely put their honey crops and relationships at risk by bottling honey that hasn’t been fully processed by bees to a net water percentage under 19%. Watery honey both feels weird and is prone to unintended fermentation. Choosing properly capped frames goes a long way to eliminating this problem. If you live in a humid place like Maryland, there is also some risk that open honey will dehumidify some of the local air, pushing water content back above dangerous levels. Truly dry honey can be achieved by technique and awareness, but if you are curious and want to directly assess the water content of your crop, Hanna Bäckmo gives a nice review of the styles and costs of refractometers used by beekeepers in this magazine (https://www.beeculture.com/refractometer/). Certainly, steady honey producers would benefit from investing in, and calibrating, these things.

A bit out of reach for most of us, but essential for the industry, are lab-based assays aimed at confirming honey purity. The methods used for this continue to improve, putting clumsy or sneaky honey producers on notice. Notably, honey yields can be stretched by a variety of refined or expelled sugars. This might be inadvertent, when syrup fed by beekeepers in the Fall for Winter survival lingers, capped until Spring. There is no easy answer to this, certainly not from me, but step one is to get bees through Winter safely, and then assess any remaining capped stores to see if these stores are bona fide honey or syrup that bees dried down but didn’t gobble up as it came in. Ask a beekeeper near you for help.

Photo by Meggyn Pomerleau on Unsplash

More insidiously, producers or packers might outright add less expensive fillers to their honey, increasing yields but losing some of the magic of honey. The technology used to detect such adulteration is improving, and several techniques are now used by regulators, producers and packers to make sure honey is pure. The International Honey Commission described forensic methods for honey purity nearly 30 years ago and updated these methods in 2009 (https://www.bee-hexagon.net/english/network/publications-by-the-ihc/). The U.S. Food and Drug Administration, keeping honest folks honest across the industry, regularly tests new methods against imported and domestic honey to identify so-called ‘economically motivated adulteration’. Using a well-established technique, Stable Carbon Isotope Ratio Analysis (SCIRA), the FDA recently screened bulk and bottled honey samples from eight countries whose honey is imported into the U.S. (https://www.fda.gov/food/economically-motivated-adulteration-food-fraud/fy2122-sample-collection-and-analysis-imported-honey-economically-motivated-adulteration). This test distinguishes ‘C4’ plant sources (largely grasses and grains) from ‘C3’ sources (all the plants with prettier, bee-visited, nectar-rich flowers). The test simply asks if the unexpected C4-sugars, often from corn syrup or sugar cane, are over-represented in honey. There is some tolerance of these C4 sugars due to bee management or assay imprecision but that level is quite low, maybe 7% by volume. Each country in the FDA screen had at least one suspicious honey batch, but the overall frequency of such batches was 10%, a level roughly similar to a much larger recent study in Europe and indicative that honey, by and large, is as advertised.

There are several newer techniques in play now for the high-stakes race between regulators and those who might diminish the reputation of honey. Dilpreet Singh Brar and colleagues in A comprehensive review on unethical honey: Validation by emerging techniques (Food Control 2023, 145, 109482, https://doi.org/10.1016/j.foodcont.2022.109482) describe nearly 50 ways to test your clover. Within the alphabet soup of available methods, they reveal six chromatographic platforms (basically methods to separate parts of a whole by size, electric charge or affinity to some sort of ‘bait’) with increasing sophistication. These machines should put fear in anyone whose honey is not perfectly sound.

As a geneticist, I am fascinated with so-called environmental DNA (eDNA) screens, whereby a complex soup is scrutinized for the genomes of the diverse organisms floating in it. Many will remember the application of eDNA screens worldwide to identify levels and variants of the SARS-Cov-2 virus in city and town wastewater systems (poor interns!; https://www.nih.gov/news-events/nih-research-matters/tracking-sars-cov-2-variants-wastewater). This same methodology is now widely used to confirm the botanical sources of honey, the genotypes of the bees collecting that honey and the myriad of other organisms from the hive environment. Practically, this method also precisely identifies any honey contaminant with a biological source, from corn syrup to diverse flower sources mixed in accidentally in coveted monofloral honeys. It is also a sensitive assay for honey bee disease agents.

For the past 20 years, genetic analyses of honey from hives have been used to confirm the presence of the bacterium responsible for American Foulbrood, Paenibacillus larvae. Federico Lauro and colleagues in Rapid detection of Paenibacillus larvae from honey and hive samples with a novel nested PCR protocol (International Journal of Food Microbiology 2003, 81, 195-201, https://doi.org/10.1016/S0168-1605(02)00257-X) showed the value of this technique for keeping track of non-symptomatic P. larvae populations. More broadly, Leigh Boardman and others have confirmed that this technique can provide a snapshot of the whole range of microbes found in colonies (Boardman, L., P. Marcelino, J. A., Valentin, R. E., Boncristiani, H., Standley, J. M., & Ellis, J. D. Novel eDNA approaches to monitor Western honey bee (Apis mellifera L.) microbial and arthropod communities. Environmental DNA. 2023; https://doi.org/10.1002/edn3.419). Here, colony-collected honey is analogous to the worker-bee samples now used in many disease surveys. Honey collections have the added value of pointing out long-ago arrivals, providing a sort of fossil record for the plants and other organisms a colony might have come into contact with during the past year. The genetic methods behind these screens are astoundingly sensitive (remember, viruses floating alone in tons of sewer sludge) and honey or hive-based screens have promise for anything from virus outbreaks to the detection of newly invasive mites and other pests. It is incredibly hard to pass through an environment without shedding a little DNA, and a little goes a long way for these sensitive methods.

Economically motivated adulteration is detectable with some effort and that’s a good thing for all of us. Honey screening, especially with the twist of identifying genetic signals from hive organisms, is also becoming a nice tool for scientists keen on monitoring disease, plant sources and the genes of the bees that did all the work.

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Found in Translation

Gut Microbes Help Bees Survive the Season
By: Jay Evans, USDA Beltsville Bee Lab

It will surprise most Bee Culture readers that microbes come in flavors that can be good, bad or indifferent to the health of their honey bee hosts. As we approach Fall, it is tempting to focus on the microbes on the good side and try to find out how to feed them for bee health prior to Winter. As someone who studies honey bee disease, I can’t help but focus on the good microbes that might interfere with agents of harm lurking in our beehives.

Kirk Anderson and colleagues in the USDA’s Tucson Carl Hayden Bee Research Laboratory have been exploring the impacts of gut microbes on bee health for a decade now. In past work, they showed how these microbes are beneficial in the guts of bees but generally ‘don’t’ help in the processing of pollen stored as bee bread. They have also shown how queens and workers differ greatly in the microbes they harbor and the impacts of bee contact on moving microbes around (see Anderson’s ‘Google Scholar’ profile for lists of his papers on these topics; https://scholar.google.com/citations?user=JiEFFkIAAAAJ&hl=en&oi=ao).

They have also explored how bees suffer mortality when the delicate microbial balance is upset. Recently, they have investigated honey bee overwintering, testing for the right mixes of nutrition and temperature that improve the odds of colony survival (hint: cold is good, to a certain degree). In a paper this past year, they describe how the gut microbes of bees react before and during Winter, building the case that microbes are critical for overwintering success (Anderson, K.E.; Maes, P. Social microbiota and social gland gene expression of worker honey bees by age and climate. Scientific Reports 2022, 12, 10690, doi:10.1038/s41598-022-14442-0).

They also show that the overwintering environment can favor certain microbes that are less helpful for bee health. Specifically, bee colonies overwintered in a warm environment started with the typical population of gut bacteria but that population broke bad in the end, notably thanks to overgrowth (these were NOT found in bees) as well as several types which ARE known to decrease bee health. Just what it is about warmer Winter environments that favors an odd, and apparently harmful, bacterial group is not known, and solving this will be key in future work aimed at prepping bees for current or future Winter climates.

More generally, disease agents are opportunists; taking advantage of their victims when something else is out of whack. These opportunities can arise from stressors in the environment, poor genetics or inadequate nutrition. Opportunities might also arise when populations of good bacteria are somehow absent. There are a myriad of ways that such ‘good’ microbes could help bees in the face of disease, from providing a physical layer on the gut wall that frustrates pathogens, to improving nutrient transfer or stimulating bee immunity.

Finally, gut microbes might directly attack the bad actors. Studies showing increased honey bee disease following heavy antibiotic treatments provide ample evidence for the roles of natural bee bacteria. In one such study, led by Jiang Hong Li and my USDA colleague Judy Chen (Li, J.H.; Evans, J.D.; Li, W.F.; Zhao, Y.Z.; DeGrandi-Hoffman, G.; Huang, S.K.; Li, Z.G.; Hamilton, M.; Chen, Y.P. New evidence showing that the destruction of gut bacteria by antibiotic treatment could increase the honey bee’s vulnerability to Nosema infection. PloS one 2017, 12, e0187505, doi:10.1371/journal.pone.0187505). Gut microbes were shown to help bees resist nosema disease. A cleansing of gut bacteria by an intensive antibiotic regime resulted in shorter lifespans overall, and increased the impacts of nosema exposure on longevity.

Sean Leonard and colleagues, in the University of Texas laboratory of Nancy Moran, showed that a human assist can further sharpen the impacts of natural gut microbes on bee parasites. Specifically, they engineered (in the laboratory) a common ‘good’ bacterium of bees so that it targeted challenges as distinct as Varroa mites and Deformed wing virus (Leonard, S.P.; Powell, J.E.; Perutka, J.; Geng, P.; Heckmann, L.C.; Horak, R.D.; Davies, B.W.; Ellington, A.D.; Barrick, J.E.; Moran, N.A. Engineered symbionts activate honey bee immunity and limit pathogens. Science 2020, 367, 573-576, doi:10.1126/science.aax9039).

Nosema. Credit: Qiang Huang

Work this year based on the same strategy (led by Qiang Huang from Jiangxi University, working in Moran’s lab) showed resident bacteria could be altered to successfully target nosema disease (Huang, Q.; Lariviere, P.J.; Powell, J.E.; Moran, N.A. Engineered gut symbiont inhibits microsporidian parasite and improves honey bee survival. Proceedings of the National Academy of Sciences 2023, 120, e2220922120, doi:10.1073/pnas.2220922120). Bees with the engineered bacteria both lived significantly longer and had far fewer nosema spores to pass on to their nestmates. Interestingly, bees fed the gut bacterium alone, and the bacterium with a nonspecific (not targeting nosema), modification also showed signs of reducing disease impacts, supporting the evidence that the bacterium itself is also a friend to bees.

Short of this high-tech solution, are there ways that beekeepers can help nurture the natural gut bacteria found in their beehives? If you supplement your bees, a recent paper by Elijah Powell and others in Moran’s group suggests that pollen-based supplements tend to lead to a more balanced ‘core’ set of bacteria in the bee gut, possibly decreasing the threats from at least one bacterial pathogen of adult bees (Powell JE, Lau P, Rangel J, Arnott R, De Jong T, Moran NA (2023) The microbiome and gene expression of honey bee workers are affected by a diet containing pollen substitutes. PLoS ONE 18(5): e0286070. https://doi.org/10.1371/journal.pone.0286070). I know there are many colony supplements available and I don’t claim this makes a pollen-based supplement better for bees overall than supplements with a different protein source (nor, of course, does this represent any formal endorsement of one type of bee feed over another). Still, it is interesting to contemplate how particular supplements affect not just bees but the hitchhiking microbes that have adapted to life in their guts.

One thing is clear from these diverse studies. While many of us focus on the microbes whose effects are damaging to bee colonies, most hive microbes are neutral or even beneficial to their bee hosts in Summer and Winter. Bees have been harnessing this power for millennia, and we would do well to help them sustain the right mix of gut partners.

Dr. Tracy Farone

Technical Updates
By: Dr. Tracy Farone

It is mid-May here in the foothills of Pennsylvania. The locust trees are in full bloom. It looks like it will be a good year for them. “Good for the bees,” says the beekeeper voice in my head. The white-tailed deer have changed color into that beautiful reddish brown that pops out within the fresh, green backdrop of the woods. As my “barn” cat (but not really a barn cat), Sylvester, snoozes, stretched out at my feet, I just watched a doe trot away from a salt block 20 yards from my deck. I am a couple of days out from the end of the semester, time to take a breath…The last thing I want to think about is meetings, committees and the possible political acrobatics that go along with them.

I must admit I usually really hate meetings… “analysis paralysis,” pre-determined “communication,” hours of my life I will never get back, things “old” people do, and such. I have always thought it ironically funny that “committee” is the term for a gathering of vultures. But I am also appreciating the importance of voicing and hearing different perspectives on issues and how it’s extremely important in today’s world. And those that step up and serve on organizational committees are giving up their valuable time to contribute to important and ever on-going work.

As promised, I would like to give you an update and summary on a few exciting collaborations that have recently taken place and hopefully bring about positive relationships and outcomes between the beekeeping industry and veterinarians. The American Veterinary Medical Association’s (AVMA) Animal Agriculture Liaison Committee (AALC) Meeting was held at AVMA Headquarters in Schaumburg, IL May 3-4, 2023. I had the opportunity to be a “fly on the wall” at times as an alternate delegate via ZOOM for some of the meeting. The Honey Bee Health Coalition’s (HBHC) Annual Meeting in Sacramento, CA was held at the same time. Both meetings hosted veterinarians representing honey bee medicine for the FIRST time. All representatives were veterinarians also serving on the Honey Bee Veterinary Consortium (HBVC) board.

The American Veterinary Medical Association’s (AVMA) Animal Agriculture Liaison Committee (AALC) Meeting Summary:
I have been an alternate delegate representing honey bees on this committee for four to five months now. I am still trying to figure out the ropes, doing mostly listening (a benefit to being the alternate). I can say the committee is continually active with legislative consulting and policy considerations coming to my email box every other day. I can also say that the committee is absolutely enthralled to learn more about honey bees. As an alternate, I did not attend the meeting in person, but Dr. Terri Kane was there, near Chicago, representing. I jumped into the meeting via ZOOM when I could. Some other perspectives include those that represent veterinarians and producers in the areas of veterinary pharmacology, bovine, fish, aquatics, swine, small ruminants, sheep, public health, cattle, chickens, turkeys and the reproduction of animals, as well as government entities like the FDA and USDA.

Discussions include topics like, the Farm Bill; various drug regulation bills; protective measures for maintaining a safe food supply; humane guidelines in animal handling; policies for identifying, preventing, and controlling several current disease threats; and reports on current issues affecting each industry represented and any on-going actions in place. Our honey bee report included information on the progress made within the HBVC and multiple Colleges of Veterinary Medicine to increase honey bee related education of veterinarians and veterinary students to better serve the industry through grant projects, additional curriculum and certification programs for practicing veterinarians. I wish I could get into more detail, but I am bound by a non-disclosure agreement and a secret handshake (just kidding about the handshake). Maybe I will work on the handshake when I attend a meeting in the flesh.

The Honey Bee Health Coalition’s (HBHC) Annual Meeting Summary:
The stated purpose of the HBHC annual meeting is to “advance dialogue and action across workstreams in the priority areas of forage and nutrition, hive management and crop pest control.” Focuses included almond production, bee protection, The Bee Integrated Demonstration Project and building relationships within members. Drs. Kristol Stenstrom and Britteny Kyle represented veterinarians and the HBVC, a new member of the HBHC, again for the first time. Various reports were shared on the status of honey bees, pollinators and the industry from both the agricultural and conservational perspectives. Best practices and projects involving disease management, habitat management and pesticide use were working topics of discussion.

Next on the List: Euthanasia and Depopulation Procedures in Honey Bees.
The AVMA is extremely interested in learning more about recommendations and guidelines for euthanizing honey bee colonies in various situations, in the safest and most humane manner. Various situations include smaller verses larger operations, stationary hives, migratory hives, emergency de-population procedures, euthanasia for public safety reasons and euthanasia for disease mitigation reasons. AVMA recommendations and guidelines exist for nearly every type of animal that veterinarians work with, except honey bees. I have been asked to be part of a special sub-committee to consider, write up and present recommendations and guidelines to the AVMA. As we begin this work, I am open to reader’s suggestions on the topic. Oh boy… another committee, here we go!