DOI: 10.5061/dryad.rjdfn2zpw
title: Substantial variation in larval honey bee nutrition within and among <em>Apis mellifera</em> colonies

description: Parents have evolved strategies to reduce the risk of malnutrition in
 offspring, including the production  of specialized nutritional
 secretions that are tailored to meet the unique needs of developing
 offspring. Studies in vertebrates, however, show abundant individual
 variation in nutritional secretions; the causes and consequences of this
 variation, and the extent to which such patterns can be generalized beyond
 vertebrates, remain unclear. Here, we investigated natural variation in
 nutritional secretions in an invertebrate species, the Western honey bee
 (Apis mellifera L.). This is a unique bee wherein developing larvae
 subsist entirely on “jellies” (e.g., royal jelly) produced by glands in
 adult worker bees. We assess within- and among-colony variation in the
 macronutrient content of the secretions fed to female worker larvae
 (“worker jelly”), collected from individual larval honeycomb cells.
 Although female workers make up the largest demographic inside a honey bee
 colony, very few studies have investigated their larval diet; even fewer
 have included the scope of colonies needed to assess natural variation in
 this critical nutritional substance. In one of the largest such studies to
 date, we found significant variation both within and among colonies in
 total quantity and macronutrient content of worker jelly, but with greater
 variation among colonies; this pattern was strongest for proteins. We
 further assessed whether worker jelly composition was correlated with
 colony defensive aggression because of extensive links between aggression,
 foraging activity, and larval development outcomes; however, we observed
 no such relationship. This study is a critical step in understanding the
 evolution and maintenance of offspring provisioning strategies, as well as
 bee foraging ecology and nutritional stress response.
description: # Substantial variation in larval honey bee nutrition within and among
 *Apis mellifera* colonies Dataset DOI:
 [10.5061/dryad.rjdfn2zpw](10.5061/dryad.rjdfn2zpw) ## Description of the
 data and file structure These data were used to generate the paper titled,
 "Substantial variation in larval honey bee nutrition within and among
 *Apis mellifera* colonies." Detailed methods can be found in the body
 of the manuscript. ### Files and variables #### File:
 WorkerJelly2025_Data.csv **Description:** This is the primary dataset that
 was used to generate all statistical analyses and figures in the main body
 of the manuscript, as well as most of the supplemental figures. #####
 Variables * Hive: A unique identifier of the colony the worker jelly was
 sourced from * Colony: A separate unique identifier of the colony that
 specifies colony aggression level and is ordered by sample collection date
 * ExtractionDate: The calendar date the worker jelly sample was collected
 * Day: Notes the day within the entire experimental window that the sample
 was collected, with "1" being the first extraction day and
 "23" being the final extraction day * UniqueName: A unique
 identifier of the sample; the first portion is the colony name while the
 second portion is the cell number (with cell numbers reused between days)
 * Aggression: "Colony Aggression," the aggression level of the
 source colony; either "high" or "low" * AggScore: The
 raw aggression score of the colony * AggRank: The rank of the aggression
 score of the colony * AggRank_Centered: Similar to above--the rank of the
 aggression score of the colony, with all colonies centered around zero *
 Source: Genetic strain or purchasing source of the colony the worker jelly
 was sourced from; "Wild stock" is Open-mated * Location:
 Identifies the yard the colony was located in, also called
 "Site"; "UpperLower" is Alpha, "Outer" is
 Beta, and "Doug" is Gamma * TotalDryWeight: The total dry mass
 of the sample (mg) * MassCorrectedProtein: The relative amount of proteins
 in the sample, corrected for the total dry weight of the sample
 (TotalProtein/TotalDryWeight) * TotalProtein: The total amount of proteins
 in the sample (mg) * MassCorrectedLipids: The relative amount of lipids in
 the sample, corrected for the total dry weight of the sample
 (TotalLipids/TotalDryWeight) * TotalLipids: The total amount of lipids in
 the sample (mg) * MassCorrectedCarbs: The relative amount of carbohydrates
 in the sample, corrected for the total dry weight of the sample
 (TotalCarbs/TotalDryWeight) * TotalCarbs: The total amount of
 carbohydrates in the sample (mg) #### File: BFCV.csv **Description:** This
 is a secondary dataset. It displays the coefficients of variation for each
 macronutrient. It was only used to create one supplemental figure (Fig.
 S1). ##### Variables * Colony: A unique identifier of the colony that
 specifies colony aggression level and is ordered by sample collection date
 * Carbohydrates: The coefficients of variation (standard deviation/mean, a
 unitless value) for all carbohydrate total mass measurements for a given
 colony. * Lipids: The coefficients of variation (standard deviation/mean,
 a unitless value) for all lipid total mass measurements for a given
 colony. * Proteins: The coefficients of variation (standard
 deviation/mean, a unitless value) for all protein total mass measurements
 for a given colony. #### File: BF_Ranges.csv **Description:** This is a
 secondary dataset. It displays the minimum and maximum value for each
 macronutrient. It was only used to create one supplemental figure (Fig.
 S2). ##### Variables * Hive: A unique identifier of the colony the worker
 jelly was sourced from * Aggression: "Colony Aggression," the
 aggression level of the source colony; either "high" or
 "low" * ProteinsMin: The minimum recorded protein value
 ("TotalProtein", in mg) for a given colony. * ProteinsMax: The
 maximum recorded protein value ("TotalProtein", in mg) for a
 given colony. * LipidsMin: The minimum recorded lipid value
 ("TotalLipids", in mg) for a given colony. * LipidsMax: The
 maximum recorded lipid value ("TotalLipids", in mg) for a given
 colony. * CarbsMin: The minimum recorded carbohydrate value
 ("TotalCarbs", in mg) for a given colony. * CarbsMax: The
 maximum recorded carbohydrate value ("TotalCarbs", in mg) for a
 given colony. #### File: ColonyAggression.csv **Description:** This is a
 secondary dataset. It condenses the colony aggression score data and
 averages the nutrient information for that colony. It was only used to
 generate two supplemental figures (Fig. S3 and S8). * Hive: A unique
 identifier of the colony the worker jelly was sourced from *
 ExtractionDate: The calendar date the worker jelly sample was collected *
 Day: Notes the day within the entire experimental window that the sample
 was collected, with "1" being the first extraction day and
 "23" being the final extraction day * Aggression: "Colony
 Aggression," the aggression level of the source colony; either
 "high" or "low" * AggScore: The raw aggression score
 of the colony * AggRank: The rank of the aggression score of the colony *
 AggRank_Centered: Similar to above--the rank of the aggression score of
 the colony, with all colonies centered around zero * Source: Genetic
 strain or purchasing source of the colony the worker jelly was sourced
 from; "Wild stock" is Open-mated * Location: Identifies the yard
 the colony was located in, also called "Site";
 "UpperLower" is Alpha, "Outer" is Beta, and
 "Doug" is Gamma * TotalDryWeight: The average total dry mass of
 a sample (mg), averaged across all samples for a given colony. *
 MassCorrectedProtein: The relative amount of proteins in each sample,
 corrected for the total dry mass of the sample
 (TotalProtein/TotalDryWeight), averaged across all samples for a given
 colony. * MassCorrectedLipids: The relative amount of lipids in each
 sample, corrected for the total dry mass of the sample
 (TotalLipids/TotalDryWeight), averaged across all samples for a given
 colony. * MassCorrectedCarbs: The relative amount of carbohydrates in each
 sample, corrected for the total dry mass of the sample
 (TotalCarbs/TotalDryWeight), averaged across all samples for a given
 colony. 
 
 #### File: WorkerJelly2025_Code.html **Description:** HTML of the R
 markdown file of all code used for this study. #### File: S14_Code.pdf
 **Description:** PDF version of the R markdown file of all code used for
 this study. 
 
 ## Code/software Data files are in .csv format and can be viewed with any
 standard text or spreadsheet editor. All code was run in R version 4.1.2
 ## Access information Other publicly accessible locations of the data: *
 Data are also available in the supplemental materials of the linked
 manuscript ("Substantial variation in larval honey bee nutrition
 within and among *Apis mellifera* colonies.") Data was derived from
 the following sources: * N/A