Humming , creating the "humming" sound from its rapid wingbeats, while
chirping by vocalization Hummingbirds are named for the prominent humming sound their wingbeats make while flying and hovering to feed or interact with other hummingbirds. Humming serves communication purposes by alerting other birds of the arrival of a fellow forager or potential mate. The humming sound of hummingbirds is unique among flying animals, compared to the whine of
mosquitoes, buzz of
bees, and "whoosh" of larger birds.
Visual system Hummingbirds have visual properties typical of species that are both
predatory and
prey, a combination that enables them to capture insects and to hover at flowers for feeding, while simultaneously avoiding potential predators.
Anatomy Hummingbirds have an upper and lower
eyelid, and a
nictitating membrane as a translucent "third" eyelid. The
iris of hummingbird eyes is dark brown, with a black
pupil measuring 216
micrometres in diameter, and a wide, thin
cornea. The prevalence of natural eye disease in a sample of Anna's and black-chinned hummingbirds was 2.3%. The enlargement of the brain region responsible for visual processing indicates an enhanced ability for perception and processing of fast-moving visual stimuli encountered during rapid forward flight, insect foraging, competitive interactions, and high-speed courtship. Hummingbirds are highly sensitive to stimuli in their visual fields, responding to even minimal motion in any direction by reorienting themselves in midflight. While in complex and dynamic natural environments, their visual sensitivity allows them to precisely hover in place The fourth color cone would extend the range of visible colors for hummingbirds to perceive ultraviolet light and color combinations of feathers and gorgets, colorful plants, and other objects in their environment, enabling detection of as many as five non-spectral colors, including purple, ultraviolet-red, ultraviolet-green, ultraviolet-yellow, and ultraviolet-purple. Many hummingbird species exhibit a diverse vocal repertoire of chirps, squeaks, whistles and buzzes. Vocalizations vary in complexity and spectral content during social interactions, foraging, territorial defense, courtship, and mother-nestling communication. It also produces
ultrasonic vocalizations which do not function in communication. The avian vocal organ, the
syrinx, plays an important role in understanding hummingbird song production. What makes the hummingbird's syrinx different from that of other birds in the Apodiformes order is the presence of internal muscle structure, accessory cartilages, and a large
tympanum that serves as an attachment point for external muscles, all of which are adaptations thought to be responsible for the hummingbird's increased ability in pitch control and large frequency range. Hummingbird songs originate from at least seven specialized
nuclei in the
forebrain. A
genetic expression study showed that these nuclei enable
vocal learning (ability to acquire vocalizations through imitation), a rare trait known to occur in only two other groups of birds (
parrots and
songbirds) and a few groups of mammals (including humans,
whales and dolphins, and
bats). Generally, birds have been assessed to vocalize and hear in the range of 2–5 kHz, with hearing sensitivity falling with higher frequencies. During flight and hovering, oxygen consumption per gram of muscle tissue in a hummingbird is about 10 times higher than that measured in elite human athletes. Hummingbirds are rare among vertebrates in their ability to rapidly make use of ingested sugars to fuel energetically expensive hovering flight, powering up to 100% of their metabolic needs with the sugars they drink. Hummingbird flight muscles have extremely high capacities for
oxidizing carbohydrates and
fatty acids via
hexokinase,
carnitine palmitoyltransferase, and
citrate synthase enzymes at rates that are the highest known for vertebrate
skeletal muscle. To sustain rapid wingbeats during flight and hovering, hummingbirds expend the
human equivalent of 150,000
calories per day, an amount estimated to be 10 times the energy consumption by a
marathon runner in competition. Hummingbirds can use newly ingested sugars to fuel hovering flight within 30–45 minutes of consumption. These data suggest that hummingbirds are able to oxidize sugar in flight muscles at rates rapid enough to satisfy their extreme metabolic demands as indicated by a 2017 review showing that hummingbirds have in their flight muscles a mechanism for "direct oxidation" of sugars into maximal
ATP yield to support a high metabolic rate for hovering, foraging at altitude, and migrating. This adaptation occurred through the
evolutionary loss of a key
gene,
fructose-bisphosphatase 2 (
FBP2), coinciding with the onset of hovering by hummingbirds estimated by fossil evidence to be some 35 million years ago. Without
FBP2,
glycolysis and mitochondrial respiration in flight muscles are enhanced, enabling hummingbirds to metabolize sugar more efficiently for energy. The amount of fat (1–2 g) used by a migrating hummingbird to cross the Gulf of Mexico in a single flight is similar to that used by a human climbing about .
Heat dissipation The high metabolic rate of hummingbirds – especially during rapid forward flight and hovering – produces increased body heat that requires specialized mechanisms of
thermoregulation for heat dissipation, which becomes an even greater challenge in hot, humid climates. Hummingbirds dissipate heat partially by
evaporation through exhaled air, and from body structures with thin or no feather covering, such as around the eyes, shoulders, under the wings (
patagia), and feet. While hovering, hummingbirds do not benefit from the heat loss by
air convection during forward flight, except for air movement generated by their rapid wing-beat, possibly aiding convective heat loss from the extended feet. Smaller hummingbird species, such as the calliope, appear to adapt their relatively higher
surface-to-volume ratio to improve convective cooling from air movement by the wings. requires a parallel dynamic range in
kidney function. During a day of nectar consumption with a corresponding high water intake that may total five times the body weight per day, hummingbird kidneys process water via
glomerular filtration rates (GFR) in amounts proportional to water consumption, thereby avoiding
overhydration. During brief periods of water deprivation, however, such as in nighttime torpor, GFR drops to zero, preserving body water. Morphological studies on Anna's hummingbird kidneys showed adaptations of high
capillary density in close proximity to
nephrons, allowing for precise regulation of water and electrolytes.
Hemoglobin adaptation to altitude Dozens of hummingbird species live year-round in tropical mountain habitats at high altitudes, such as in the Andes over ranges of to where the
partial pressure of oxygen in the air is reduced, a condition of
hypoxic challenge for the high metabolic demands of hummingbirds. In Andean hummingbirds living at high elevations, researchers found that the oxygen-carrying protein in blood
hemoglobin had increased oxygen-
binding affinity, and that this adaptive effect likely resulted from evolutionary
mutations within the hemoglobin molecule via specific amino acid changes due to natural selection.
Adaptation to winter Anna's hummingbirds are the northernmost year-round residents of any hummingbird. Anna's hummingbirds were recorded in Alaska as early as 1971, and resident in the
Pacific Northwest since the 1960s, particularly increasing as a year-round population during the early 21st century. Scientists estimate that some Anna's hummingbirds overwinter and presumably breed at northern latitudes where food and shelter are available throughout winter, tolerating moderately cold winter temperatures. In addition, hummingbirds with inadequate stores of body fat or insufficient plumage are able to survive periods of subfreezing weather by lowering their metabolic rate and entering a state of
torpor. While their range was originally limited to the
chaparral of California and
Baja California, it expanded northward to
Oregon,
Washington, and
British Columbia, and east to
Arizona over the 1960s to 1970s. In the Pacific Northwest, the fastest growing populations occur in regions with breeding-season cold temperatures similar to those of its native range. and is a non-migrating resident of
Seattle where it lives year-round through winter enduring extended periods of subfreezing temperatures, snow, and high winds.
Torpor The metabolism of hummingbirds can slow at night or at any time when food is not readily available; the birds enter a deep-sleep state (known as torpor) to prevent energy reserves from falling to a critical level. with heart and
breathing rates slowing dramatically (heart rate of roughly 50 to 180 bpm from its daytime rate of higher than 1000 bpm). Recordings from a
Metallura phoebe hummingbird in noctural torpor at around in the Andes mountains showed that body temperature fell to 3.3 °C (38 °F), the lowest known level for a bird or non-hibernating mammal. During cold nights at altitude, hummingbirds were in torpor for 2–13 hours depending on species, with cooling occurring at the rate of 0.6 °C per minute and rewarming at 1–1.5 °C per minute. Use and duration of torpor vary among hummingbird species and are affected by whether a dominant bird defends territory, with nonterritorial subordinate birds having longer periods of torpor. A hummingbird with a higher fat percentage will be less likely to enter a state of torpor compared to one with less fat, as a bird can use the energy from its fat stores. Among the better-known North American species, the typical lifespan is probably 3 to 5 years. The longest recorded lifespan in the wild relates to a female broad-tailed hummingbird that was banded as an adult at least one year old, then recaptured 11 years later, making her at least 12 years old. Other longevity records for banded hummingbirds include an estimated minimum age of 10 years 1 month for a female black-chinned hummingbird similar in size to the broad-tailed hummingbird, and at least 11 years 2 months for a much larger
buff-bellied hummingbird. ==Natural enemies==