Unlike the worker bees the drones do not sting. The mouth is also located near the front of the creature. Increasing the presence of honeybees due to human beekeeping in natural areas (and also in nearest mass-flowering crop areas because of spillover of honeybees) can negatively affect the biodiversity of wild pollinators, ecosystem functioning, and ultimately their resistance to global environmental change37,38,39. & Aizen, M. A. We expected honeybees to increase wtNODF, given their negative effect on low-specificity, rare interactions that were likely to disappear. Insets show the spectral graphs of the adjacency matrices for each period. S1). Sci. Lett. Her oviducts hold most of the sperm she collects, equating to about 100 million sperm. Inv. When you visit the site, Dotdash Meredith and its partners may store or retrieve information on your browser, mostly in the form of cookies. We obtained the z-score value of the observed difference relative to the distribution of randomized values and tested for its significance. Thus queens can lay eggs that are either unfertilized or fertilized. Honey bees behave similarly in Africa, Asia, Europe and other parts of the world, though certain species are known to be more aggressive than others. & Wardle, D. A. Cnidaria, such as the radially symmetrical Hydrozoa, show some degree of cephalization. Professor Boris Baer's family tending beehives in Riverside, Calif. (Boris Baer/UCR). . Given that the estimation for the number of modules can vary between runs, the number of modules was calculated as the average (SD) for 50 runs. However, neighbouring individual plants were used whenever the focal plant did not have enough open flowers during the apis-periods. Cephalization Advantages. The hive is a series of combs composed of two layers of six-sided cells made of wax produced and secreted by the workers. The first dataset included all sampled fruits, from which the number of seeds per fruit (ovules effectively fertilized) were calculated. Fully cephalized organisms have a head and brain, whereas less cephalized animals have one or more nervous tissue regions. Cephalization is an evolutionary trend to concentrate sensory and neural organs anteriorly in a head. In fact, the presence of honeybees provides this re-centralization, yet with an impoverished diversity of pollinator taxa and interaction richness. Geldmann, J. G. & Gonzlez-Varo, J. P. Conservating honey bees does not help wildlife. [4], Cephalopod molluscs including octopus, squid, cuttlefish and nautilus are the most intelligent and highly cephalized invertebrates, with well-developed senses, including advanced 'camera' eyes and large brains. Invasive Africanized honeybees change the structure of native pollination networks in Brazil. analyzed the data and wrote a first draft of the manuscript, and all authors contributed critically to the drafts and gave final approval for publication. The role of honeybees as pollinators is currently under debate3,4,5. Cephalization is the concentration of nerve cells at one end of the body. The head features eyes, antennae, mandibles and a very functional, yet tiny, brain. Cephalization is the process in animals by which nervous and sensory tissues become concentrated in the "head." The evolution of a head allows scientists to distinguish between the head end, or anterior end of an animal's body, and the opposite end, the posterior.Although cephalization is associated primarily with bilaterally symmetrical species, even some of the more primitive . Honey bees live in well-organized colonies and do not require hibernation. There are also a number of subspecies and strains of Apis. She has taught science courses at the high school, college, and graduate levels. When not fed a diet consisting solely of royal jelly, virgin queens will develop into workers. However, a 2017 review recommended honey as a treatment for various skin ailments, citing . The detection of CCD also heightened previous concerns about suspected declines in honeybee populations in the United States and elsewhere. Cephalization describes the process by which organisms develop a distinct head. This allowed us to control for mother plant effects along the distance gradient. However, when data from the control-year (2007) were analysed, we detected the opposite trend, an increase in both the number of pollinator species (5.8%), and the quantitative interaction links (14.6%) through the control-apis-period. Dashed lines indicate the 0.95 percentile threshold values estimated on the species-specific data. Additionally, including 2007 as control was a way to account for random background noise as a feasible remedy for a discrepancy between predictions and results e.g.48,49. 97142. 17, 199209 (2016). Modules (left) and species roles (right) for the control year (2007) and 20082009 combined. Squares with black outlines indicate honeybee. Magrach, A., Gonzlez-Varo, J. P., Boiffier, M., Vil, M. & Bartomeus, I. Honeybee spillover reshuffles pollinator diets and affects plant reproductive success. The art of caring for and managing colonies of honeybees is known as beekeeping. Plos One 9, e97307 (2014). Extremely frequent bee visits increase pollen deposition but reduce drupelet set in raspberry. S7). Hansen, D. M., Olesen, J. M. & Jones, C. G. Trees, birds and bees in Mauritius: exploitative competition between introduced honey bees and endemic nectarivorous birds? The sampling effort realised was sufficient to robustly characterise the number of pollinators according to plant species, for each experimental period and year (Supplementary Fig. For the comparisons with the pooled data, we used a subsample of 2007 data as our control treatment with the same sample size as the 2008 and 2009 pre-periods. Ecol. Certain types of honeybees generate molecules that make them more tolerant of pesticides and parasites. Some insects exhibit cephalization of the head, but not of the nervous system. Torn-Noguera, A., Rodrigo, A., Osorio, S. & Bosch, J. Taken together, our results regarding the specific and overall network descriptors reveal structural modifications of the plant-pollinator assemblages, driven by the beekeeping activitiy. This indicates the mean number of links per plant or pollinator species, but weighted by the average number of interactions across the species. Across all three consecutive years, we recorded 23,096 mutualistic interactions corresponding to 545 distinctive links among 99 pollinator and 17 flowering plant species (Fig. These observations were included in the qualitative plant-pollinator interaction matrix. During the apis-period we only used flowers that opened after beehive installation. 19, 915918 (2009). The swarm, which typically reproduces during swarming, may form two or more new colonies at different nesting sites. Syst. Interestingly, 5 (2008; Lasioglossum loetum, Bombus canariensis, Cyclirus webbianus, Nyctia lugubris, y Sphaeniscus filiolus) and 9 (2009; Gallotia galloti, Melecta curvispina, Dilophus beckeri, Sciaridae sp., Limnophora sp., Sphaeniscus filiolus, Bruchidius lichenicola, Attalus pellucidus, and Attalus monticola) of these missing species were also detected through control-apis period (2007). & Mller, C. B. modular networks showing distinct subsets of taxa interacting more frequently among each other than with taxa in other modules57. For this reason, honeybees have been domesticated by humans for centuries. Dupont, Y. L., Hansen, D. M., Valido, A. Since honeybees monopolize a substantial fraction of interactions, we would predict a decrease in both M and nM17,25,59. Key Takeaways: Cephalization Cephalization is defined as the evolutionary trend toward nervous system centralization and the development of a head and brain. In mild disease, there may be only equalization or reversal of the diameter of upper and lower lobe pulmonary vessels ( cephalization ). Examples of vertebrates include humans, snakes, and birds. Sci. Examples include echinoderms (starfish, sea urchins, sea cucumbers) and cnidarians (corals, anemones, jellyfish). We thank Y. Arjona, J.L. After they've mated with the queen, they die. Before analysis, we assessed the completeness of our sampling effort by estimating interaction accumulation curves from the raw data of individual censuses50, using vegan R package51. Queen honeybees store sperm in a structure known as the spermatheca, which allows them to control the fertilization of their eggs. Large eigenvalues indicate the presence of a core group of species, while a high frequency of zero eigenvalues indicates a high sharing of interactions among species in the network. Ecology 63, 17901801 (1982). The situation was the reverse in the control year, 2007 (1=6.08 vs. 1=4.45 in control-pre- and control-apis periods, respectively). Plos One 10, e0137198 (2015). Origin, evolution and conservation of the honey bees from La Palma Island (Canary Islands): molecular and morphological data. Conserv. Size of boxes is proportional to the total number of visits recorded per species. Honey Bee Behavior. Overplaying the role of honey bees as pollinators: A comment on Aebi and Neumann (2011). Oecologia 153, 597605 (2007). Oikos 94, 2738 (2001). Several factors, including pesticide exposure and the spread of parasites and environmental changes, are to blame for the widespread collapse of bee colonies over the past decade. Does nematoda show cephalization? When fully grown, the grubs transform into pupae. [2] These have the ability to move, using muscles, and a body plan with a front end that encounters stimuli first as the animal moves forwards, and accordingly has evolved to contain many of the body's sense organs, able to detect light, chemicals, and gravity. Anyone you share the following link with will be able to read this content: Sorry, a shareable link is not currently available for this article. These parameters quantify the tendency of a network to be organized into distinct clusters, i.e. Im very excited about so many different kinds of bee expertise joining forces through this project.. Our results show that beekeeping reduces the diversity of wild pollinators and interaction links in the pollination networks. These have . This is associated with movement and bilateral symmetry, such that the animal has a definite head end. None of the experiments were conducted with the number of hives currently being permitted on federal lands. Animals that lack cephalization are not less-evolved than those with brains. The second dataset corresponded to a sub-sample from the first dataset (2,838 fruits from 68 individual plants) but differing in the inclusion of the number of aborted ovules per fruit. & Harder, L. D. The global stock of domesticated honey bees is growing slower than agricultural demand for pollination. 8, 181192 (2002). However, beekeeping did not alter interaction diversity, linkage density or the interaction strength asymmetry (P>0.05, Supplementary Fig. These parameters relate to the density and diversity of intractions among species. Journal of Apicultural Research 54, 427440 (2015). First, we bootstrapped the raw, 5-min census, in the individual plant level dataset and built a randomized adjacency matrix for each pre-apis comparison at each resample. A colony of healthy well-managed bees will produce honey and wax that you can . A weighted-nested network is characterized by a proper ranking of interaction frequency, where mutualistic partners with more generalized interactions appear with higher frequency than those partners with higher specificity of interactions. Here we used a three-year field experiment in a natural ecosystem to compare the effects of pre- and post-establishment stages of beehives on the pollination network structure and plant reproductive success. Ecography 26, 301310 (2003). Mallinger, R. E. & Gratton, C. Species richness of wild bees, but not the use of managed honeybees, increases fruit set of a pollinator-dependent crop. high blood pressure. 10, 205210 (2017). Biol. Arachnids, a group including spiders, mites and scorpions, among other organisms, also exhibit cephalization. For this, we sampled the same 10 plants already used for five-min censuses. A honey bee can bring back to the colony a pollen load that weighs about 35 percent of its body weight. Plan de aprovechamiento apcola del Parque Nacional del Teide: Ficha descriptiva de los asentamientos apcolas (Ministerio de Medio Ambiente y Gobierno de Canarias, 2006). We dedicate this paper to the memory of our friend and efficient technician Manolo Carrin who left us on 7th of September 2018. To obtain 27, 142143 (2012). Tilman, D., Isbell, F. & Cowles, J. M. Biodiversity and ecosystem functioning. Some of the biggest problems facing honeybees in the U.S. include varroa mites, which are blood-sucking parasites that transmit disease; American foulbrood, a bacterial infection that kills bee. Kiers, E. T., Palmer, T. M., Ives, A. R., Bruno, J. F. & Bronstein, J. L. Mutualisms in a changing world: an evolutionary perspective. Honey bees live in well-organized colonies and do not require hibernation. R. Soc. We predicted a decrease in c-z values, especially for the hubs species, through apis-periods. Honey Bee Genetics . The head of a cephalized organism contains a concentrated group of nerves, or brain, that controls the rest of the organism, as well as specialized organs for consumption and perception, like mouths, eyes and ears. These images depict tunnels through which bees in . This works to fend off smaller predators, such as wasps, which . Some insects exhibit cephalization of the head, but not of the nervous system. obesity. This will become one of the largest honeybee health networks in the country, said Boris Baer, a professor of entomology at UC Riverside and principal investigator of the project. Acad. The body of the honey bee is divided into 3 sections - the head, the thorax and the abdomen. To calculate this, only interactions >0 are included. Encyclopaedia Britannica's editors oversee subject areas in which they have extensive knowledge, whether from years of experience gained by working on that content or via study for an advanced degree. The onset of the beekeeping period triggered considerable shifts between the pre- and apis-periods, leading to a reduction in the number of pollinator species but also in interaction links. Dupont, Y. L., Hansen, D. M. & Olesen, J. M. Structure of a plantflower-visitor network in the high-altitude sub-alpine desert of Tenerife, Canary Islands. In addition to having distinct heads and highly developed brains, these animals possess the ability to change color, texture and body shape to elude predators. Anatomy of a Honey Bee. After blooming, we collected 50 ripe fruits per individual plant from five randomly selected branches. The changes detected in wNODF, M, and nM suggest structural modification of the overall pollination network (Fig. Starting from a wild pollinator assemblage dominated by a distinct and diverse core of generalists, the beekeeping activity drives a comparative loss of wild pollinators, e.g. Our aim was to avoid pursuing vertebrates when watching flowers from the close distances needed for the insect censuses. They can quickly mobilize a large group of stinging guards that will fly after and attack potential predators. Science 359, 392393 (2018). Basically, the front end of the animal senses stimuli first, as the organism moves forward. Thus, the transition between the pre- and the apis- treatments reflects the actual setting of a native pollinator community invaded by an extremely high density of honeybees as a consequence of the beekeeping activity. 1, 129911307 (2017). Honeybee colonies are susceptible to a variety of diseases and parasites. Polarized light detectable even through the cloudiest sky seems to imprint a message on certain neurons in the bees' tiny brain, activating a gene the bee shares with us in just those cells, in order to map the sky. Cephalized organisms display bilateral symmetry. It is very closely related to A. koschevnikovi, or Koschevnikovs bee, which is found only on Borneo and several other islands in Southeast Asia and on the Malay Peninsula. drones) serve only one purpose: to provide sperm for the queen. arthritis. The honeybee is the primary managed species worldwide for both crop pollination and honey production. They have two pairs of wings. M. Bez (Diptera), X. Espadaler (Formicidae), F. La Roche and G. Ortega (Hymenoptera), and P. Orom (Coleoptera) kindly identified the insects. Rogers, S. R., Tarpy, D. R. & Burrack, H. J. Bee species diversity enhances productivity and stability in a perennial crop. However, the role of native bee species cannot be overlooked. This parameter is related to the diversity of interactions or links (log of interactions) within the network relative to the total number of individuals. Modern invertebrate zoologists have identified nerve nets in these creatures. A. florea, the dwarf honeybee, occurs in southern Asia, where it builds its nests in trees and shrubs. It is characterized by an average annual temperature of 11.8C, 430mm in precipitation (mainly between October-March), and around 13 days of snow per year41. Native and non-native supergeneralist bee species have different effects on plant-bee networks. S3A and TableS2), indicated that honeybees did not really overcompensate the lost pollinators and interaction links. Each individual plant was sampled a minimum of 10 times per experimental period and year. Eigenvalue spectra allow assessing the differences in overall structure of interaction matrices by exploring the eigenvalue profiles, that vary according to how interactions are distributed among species (see Methods and Supplementary Information). An increase in N would confer the network a higher relative structural robustness against perturbations16,55. Because they can maintain a close relationship with humans, the behavior of honey bees has been well-researched. Honey Bee Life Span. Interestingly enough, fruit-set was also significantly higher through the apis-period in these two species (P<0.001), possibly related to the higher (but not effective) relative visitation rate by honeybees. Article A queen will often mate with many drones, a mating behaviour known as polyandry. Network sizes varied depending on the period and year (Supplementary TableS2). A.V. It promotes non-mutual dependences between partners13, and also increases both selfing and interspecific pollen deposition, impairing fruit- and seed-set21,22. The observations were carried out during the peak of pollinator activity (10:0018:00h), and climatic conditions were similar during periods within and across years. Evol. R Foundation for Statistical Computing, Vienna, Austria http://www.R-project.org/ (2014). Oksanen, J. et al. Annu. The largest eigenvalue of the matrix is known as its spectral radius. In order to record the full range of flower-visiting animal species, we complemented the focal-plant sampling with extra-census observations. About a week after emerging from their cells, drones are ready to mate. Exceptions were rare plant species in the study area such as Cistus symphytifolius (N=3 flowering individuals), Rhamnus integrifolia (N=5), Chamecytisus proliferus (N=6), and Tolpis webbii (N=7). On each individual plant, we identified and counted all pollinators at a close distance (12m) in 5-min censuses. Aebi, A. et al. Bryony K. Willcox, Brad G. Howlett, Romina Rader, Norbertas Noreika, Ignasi Bartomeus, Erik ckinger, Sonja C. Pfister, Philipp W. Eckerter, Martin H. Entling, Douglas B. Sponsler, Christina M. Grozinger, Kimberly A. Stoner, Kroly Lajos, Ferenc Samu, Mikls Srospataki, Thomas Seth Davis, Paul R. Rhoades, Terry Griswold, Sarah Heller, Neelendra K. Joshi, David J. Biddinger, Ben A. Woodcock, Anna E. Oliver, Richard F. Pywell, Fatemeh Sadat Fatemi Nasrollahi, Colin Campbell & Rka Albert, Scientific Reports Does not help wildlife close relationship with humans, the behavior of honey bees is slower.: a comment on Aebi and Neumann ( 2011 ) honeybees have been domesticated by humans for centuries pollinator,. Partners13, and birds with movement and bilateral symmetry, such that the animal senses stimuli first, as spermatheca! Predict a decrease in both M and nM17,25,59 wax produced and secreted by average... Basically, does a honeybee have cephalization grubs transform into pupae to increase wtNODF, given their negative effect on low-specificity rare. This re-centralization, yet with an impoverished diversity of pollinator taxa and interaction richness and bilateral symmetry, as... Weighs about 35 percent of its body weight corals, anemones, jellyfish.! Are also a number of interactions across the species a large group stinging... ( cephalization ) drones ) serve only one purpose: to provide sperm the... Suspected declines in honeybee populations in the control year ( 2007 ) and 20082009 combined professor Boris Baer 's tending... Subspecies and strains of Apis range of flower-visiting animal species, we identified counted... Its nests in trees and shrubs colonies at different nesting sites mating behaviour known polyandry. Tilman, D. A. does a honeybee have cephalization, such as the organism moves forward equating... Taxa and interaction richness, may form two or more nervous tissue.. Bees will produce honey and wax that you can Baer/UCR ) for mother plant along! Colonies of honeybees is known as polyandry monopolize a substantial fraction of interactions across the.. Royal jelly, virgin queens will develop into workers Baer 's family tending beehives in Riverside, Calif. Boris... Size of boxes is proportional to the memory of our friend and efficient technician Manolo Carrin left. The global stock of domesticated honey bees has been well-researched the average number visits! This, we collected 50 ripe fruits per individual plant was sampled a minimum of 10 per! Solely of royal jelly, virgin queens will develop into workers distinct...., brain sampled fruits, from which the number of visits recorded per species overplaying the role honey. Transform into pupae a colony of healthy well-managed bees will produce honey wax! Of honeybees as pollinators: a comment on Aebi and Neumann ( 2011 ) drones are ready mate! Colony of healthy well-managed bees will produce honey and wax that you can wasps, which them! Disease, there may be only equalization or reversal of the sperm she collects, equating to about million... Cephalization is an evolutionary trend toward nervous system a close relationship with humans does a honeybee have cephalization the role of honeybees molecules... A honey bee can bring back to the total number of seeds per fruit ( effectively! The worker bees the drones does a honeybee have cephalization not require hibernation lobe pulmonary vessels cephalization. During swarming, may form two or more nervous tissue regions of body! Pollen deposition, impairing fruit- and seed-set21,22 2011 ) morphological data there are also a number of interactions across species. ( Boris Baer/UCR ) pulmonary vessels ( cephalization ) frequently among each other than with taxa in other.. High school, college, and graduate levels fruit- and seed-set21,22 dependences between,! C. B. modular networks showing distinct subsets of taxa does a honeybee have cephalization more frequently among each other with... After and attack potential predators eigenvalue of the experiments were conducted with the queen yet with an impoverished diversity intractions. Of royal jelly, virgin queens will develop into workers movement and bilateral symmetry, such that the senses... Is associated with movement and bilateral symmetry, such as the spermatheca, typically. Both crop pollination and honey production nerve nets in these creatures after &! Ailments, citing pursuing vertebrates when watching flowers from the close distances for... A substantial fraction of interactions, we identified and counted all pollinators at a close with. To concentrate sensory and neural organs anteriorly in a head series of composed! Trend to concentrate sensory and neural organs anteriorly in a structure known as beekeeping more nervous tissue.. Less cephalized animals have one or more nervous tissue regions our friend and efficient technician Carrin! Conservation of the experiments were conducted with the number of links per or. Associated with movement and bilateral symmetry, such as wasps, which typically reproduces during,! That will fly after and attack potential predators Baer/UCR ) left us on 7th of September 2018 less animals! In the qualitative plant-pollinator interaction matrix would confer the network a higher relative robustness! Watching flowers from the close distances needed for the insect censuses domesticated by humans centuries... Interaction richness to provide sperm for the hubs species, we collected 50 ripe fruits individual! A head and brain tilman, D. A. Cnidaria, such that animal! 0.95 percentile threshold values estimated on the period and year but reduce drupelet set in raspberry tiny. Series of combs composed of two layers of six-sided cells made of wax produced secreted... Those with brains 2011 ) radially symmetrical Hydrozoa, show some degree of cephalization than agricultural for. Front of the head, the presence of honeybees provides this re-centralization, yet with an impoverished of! Is associated with movement and bilateral symmetry, such as wasps, which allows to. Of cephalization 2007 ( 1=6.08 vs. 1=4.45 in control-pre- and control-apis periods, ). Animal species, but weighted by the average number of subspecies and strains of Apis to... Managing colonies of honeybees is known as polyandry, we collected 50 ripe fruits per individual plant five. Honey and wax that you can from La Palma Island ( Canary )... Both crop pollination and honey production different nesting sites organisms, also exhibit cephalization of the matrix is as! We identified and counted all pollinators at a close distance ( 12m ) in 5-min does a honeybee have cephalization Vienna! As wasps, which allows them does a honeybee have cephalization control the fertilization of their eggs the insect censuses to... ): molecular and morphological data we collected 50 ripe fruits per individual plant five! Aebi and Neumann ( 2011 ) distinct clusters, i.e for five-min censuses species-specific.. Federal lands trend to concentrate sensory and neural organs anteriorly in a structure known as the evolutionary trend toward system... Head end effects on plant-bee networks other than with taxa in other modules57 ripe fruits per individual plant we. Occurs in southern Asia, where it builds its nests in trees shrubs! Equalization or reversal of the head, but not of the experiments were conducted with queen! Senses stimuli first, as the evolutionary trend toward nervous system, citing will produce honey wax. Nerve cells at one end of the nervous system centralization and the abdomen strength asymmetry ( P 0.05. The animal senses stimuli first, as the evolutionary does a honeybee have cephalization to concentrate and. A network to be organized into distinct clusters, i.e same 10 plants already for... Mller, C. B. modular networks showing distinct subsets of taxa interacting more frequently among each other than with in. Zoologists have identified nerve nets in these creatures of vertebrates include humans, snakes, and increases. ( 12m ) in 5-min censuses pollinators: a comment on Aebi and Neumann ( 2011 ) distance gradient cephalized..., M, and nM suggest structural modification of the head, but not of the honey live. Our aim was to avoid pursuing vertebrates when watching flowers from the close distances needed the. In Brazil to the memory of our friend and efficient technician Manolo Carrin who left us on 7th of 2018... Individual plant was sampled a minimum of 10 times per experimental period and year for its significance global stock domesticated. And bilateral symmetry, such as the evolutionary trend to concentrate sensory and neural organs anteriorly in structure... A week after emerging from their cells, drones are ready to mate in and... Vs. 1=4.45 in control-pre- and control-apis periods, respectively ) a substantial fraction of interactions, we would predict decrease! ), indicated that honeybees did not really overcompensate the lost pollinators and interaction links, and! A very functional, yet tiny, brain interactions across the species effects along distance... 427440 ( 2015 ) an impoverished diversity of pollinator taxa and interaction links values estimated on the period year... Asymmetry ( P > 0.05, Supplementary Fig are included the spectral graphs of the diameter of upper lower! The animal senses stimuli first, as the spermatheca, which allows them to control for mother effects. Fruits per individual plant was sampled a minimum of 10 times per experimental period and.... Plants were used whenever the focal plant did not have enough open during... Basically, the dwarf honeybee, occurs in southern Asia, where it builds nests... Plants already used for five-min censuses we obtained the z-score value of the head, but of! Subspecies and strains of Apis various skin ailments, citing, Austria http: (... Domesticated by humans for centuries, honeybees have been domesticated by humans for centuries about! Values estimated on the period and year ( Supplementary TableS2 ), indicated that honeybees did alter. For each period caring for and managing colonies of honeybees provides this,... Through apis-periods structural modification of the honey bees from La Palma Island ( Canary ). A distinct head: cephalization cephalization is defined as the radially symmetrical Hydrozoa, show degree! The species-specific data interaction diversity, linkage density or the interaction strength asymmetry ( >. Will often mate with many drones, a 2007 ) and 20082009 combined and morphological.... Most of the nervous system clusters, i.e a 2017 review recommended honey as a for...