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46-540: Afribyx senegallus (Linnaeus, 1766) Parra senegalla Linnaeus, 1766 The African wattled lapwing ( Vanellus senegallus ), also known as the Senegal wattled plover or simply wattled lapwing , is a large lapwing , a group of largish waders in the family Charadriidae . It is a resident breeder in most of sub-Saharan Africa outside the rainforests, although it has seasonal movements. These are conspicuous and unmistakable birds . They are large brown waders with

92-524: A black crown, white forehead and large yellow facial wattles. The tail is white, tipped black, and the long legs are yellow. In flight, the upperwings have black flight feathers and brown coverts separated by a white bar. The underwings are white with black flight feathers. The African wattled lapwing has a loud peep-peep call. This species is a common breeder in wet lowland habitats, especially damp grassland. It often feeds in drier habitats, such as golf courses, picking insects and other invertebrates from

138-427: A number of different steps within the carotenoid biosynthetic pathway. Glyceraldehyde 3-phosphate and pyruvate , intermediates of photosynthesis , are converted to deoxy-D-xylulose 5-phosphate (DXP) catalyzed by DXP synthase (DXS). DXP reductoisomerase catalyzes the reduction by NADPH and subsequent rearrangement. The resulting MEP is converted to 4-(cytidine 5’-diphospho)-2-C-methyl-D-erythritol (CDP-ME) in

184-473: A selection of true lapwings and plovers would also give a good idea of charadriid wader evolution altogether. A mid- Oligocene – c.28 mya ( million years ago ) – fossil from Rupelmonde in Belgium has been assigned to Vanellus , but even if the genus were broadly defined, it is entirely unclear if the placement is correct. Its age ties in with the appearance of the first seemingly distinct Charadriinae at about

230-496: A shrill, wailing cry. The traditional terms "plover", "lapwing", and "dotterel" do not correspond exactly to current taxonomic models; thus, several of the Vanellinae are often called plovers, and one a dotterel, while a few of the "true" plovers (subfamily Charadriinae ) are known colloquially as lapwings. In general, a lapwing can be thought of as a larger plover. In Europe's Anglophone countries, lapwing refers specifically to

276-461: A single enzyme, the bacterial phytoene desaturase (CRTI) for the catalysis. Plants and cyanobacteria however utilize four enzymes for this process. The first of these enzymes is a plant-type phytoene desaturase which introduces two additional double bonds into 15-cis-phytoene by dehydrogenation and isomerizes two of its existing double bonds from trans to cis producing 9,15,9’-tri-cis-ζ-carotene. The central double bond of this tri-cis-ζ-carotene

322-407: Is a higher energy state and is essential in photoprotection. Light produces damaging species during photosynthesis, with the most damaging being reactive oxygen species (ROS). As these high energy ROS are produced in the chlorophyll the energy is transferred to the carotenoid’s polyene tail and undergoes a series of reactions in which electrons are moved between the carotenoid bonds in order to find

368-468: Is a virulence factor with an antioxidant action that helps the microbe evade death by reactive oxygen species used by the host immune system. The basic building blocks of carotenoids are isopentenyl diphosphate (IPP) and dimethylallyl diphosphate (DMAPP). These two isoprene isomers are used to create various compounds depending on the biological pathway used to synthesize the isomers. Plants are known to use two different pathways for IPP production:

414-441: Is cyclized; cyclization gives rise to carotenoid diversity, which can be distinguished based on the end groups. There can be either a beta ring or an epsilon ring, each generated by a different enzyme ( lycopene beta-cyclase [beta-LCY] or lycopene epsilon-cyclase [epsilon-LCY]). α-Carotene is produced when the all-trans lycopene first undergoes reaction with epsilon-LCY then a second reaction with beta-LCY; whereas β-carotene

460-509: Is derived from herbicides applied to soil and binds to DXP synthase. This inhibits DXP synthase, preventing synthesis of DXP and halting the MEP pathway. The use of this toxin leads to lower levels of carotenoids in plants grown in the contaminated soil. Fosmidomycin , an antibiotic , is a competitive inhibitor of DXP reductoisomerase due to its similar structure to the enzyme. Application of said antibiotic prevents reduction of DXP, again halting

506-437: Is directly related to their structure, especially the length of the conjugation. Xanthophylls are often yellow, giving their class name. Carotenoids also participate in different types of cell signaling. They are able to signal the production of abscisic acid , which regulates plant growth, seed dormancy , embryo maturation and germination , cell division and elongation, floral growth, and stress responses. The length of

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552-455: Is eliminated from the CDP-ME2P molecule. MECDP is then converted to (e)-4-hydroxy-3-methylbut-2-en-1-yl diphosphate (HMBDP) via HMBDP synthase in the presence of flavodoxin and NADPH. HMBDP is reduced to IPP in the presence of ferredoxin and NADPH by the enzyme HMBDP reductase . The last two steps involving HMBPD synthase and reductase can only occur in completely anaerobic environments. IPP

598-554: Is imperative that female birds carefully select high quality mates. Current literature supports the theory that vibrant carotenoid coloration is correlated with male quality—either though direct effects on immune function and oxidative stress, or through a connection between carotenoid metabolizing pathways and pathways for cellular respiration. It is generally considered that sexually selected traits, such as carotenoid-based coloration, evolve because they are honest signals of phenotypic and genetic quality. For instance, among males of

644-400: Is isomerized by the zeta-carotene isomerase Z-ISO and the resulting 9,9'-di-cis-ζ-carotene is dehydrogenated again via a ζ-carotene desaturase (ZDS) . This again introduces two double bonds, resulting in 7,9,7’,9’-tetra-cis-lycopene. CRTISO , a carotenoid isomerase, is needed to convert the cis -lycopene into an all-trans lycopene in the presence of reduced FAD . This all-trans lycopene

690-403: Is no correlation between consumption of foods high in carotenoids and vitamin A and the risk of Parkinson's disease . Humans and other animals are mostly incapable of synthesizing carotenoids, and must obtain them through their diet. Carotenoids are a common and often ornamental feature in animals. For example, the pink color of salmon , and the red coloring of cooked lobsters and scales of

736-513: Is produced by two reactions with beta-LCY. α- and β-Carotene are the most common carotenoids in the plant photosystems but they can still be further converted into xanthophylls by using beta-hydrolase and epsilon-hydrolase, leading to a variety of xanthophylls. It is believed that both DXS and DXR are rate-determining enzymes, allowing them to regulate carotenoid levels. This was discovered in an experiment where DXS and DXR were genetically overexpressed, leading to increased carotenoid expression in

782-414: Is then able to isomerize to DMAPP via IPP isomerase. Two GGPP molecules condense via phytoene synthase (PSY), forming the 15-cis isomer of phytoene . PSY belongs to the squalene/phytoene synthase family and is homologous to squalene synthase that takes part in steroid biosynthesis. The subsequent conversion of phytoene into all-trans- lycopene depends on the organism. Bacteria and fungi employ

828-920: The Late Pleistocene . Little is known of this rather large lapwing; it may actually belong in Vanellus . The remaining Charadrii are highset and/or chunky birds, even decidedly larger than a lot of the scolopacid waders . The evolutionary trend regarding the Charadriidae – which make up most of the diversity of the Charadrii – thus runs contrary to Cope's Rule . Genus Vanellus Genus Hoploxypterus Genus Erythrogonys Lipochrom All are derivatives of tetraterpenes , meaning that they are produced from 8 isoprene units and contain 40 carbon atoms. In general, carotenoids absorb wavelengths ranging from 400 to 550 nanometers (violet to green light). This causes

874-535: The Quaternary . The Early Oligocene fossil Dolicopterus from Ronzon, France may be such an ancestral member of the Charadriidae or even the Vanellinae, but it has not been studied in recent decades and is in dire need of review. Apart from the prehistoric Vanellus , the extinct lapwing genus Viator has been described from fossils. Its remains were found in the tar pits of Talara in Peru and it lived in

920-460: The northern lapwing , the only member of this group to occur in most of the continent and thus the first bird to go by the English name lapwing (also known as peewit or pyewipe ). In the fanciful taxonomy promoted by medieval courtesy books , a group of lapwings was called a "deceit". While authorities generally agree that there are approximately 25 species of Vanellinae, classifications within

966-513: The Vanellinae is scant and mostly recent in origin; no Neogene lapwings seem to be known. On the other hand, it appears as if early in their evolutionary history the plovers, lapwings and dotterels must have been almost one and the same, and they are hard to distinguish osteologically even today. Thus, since the Red-kneed Dotterel is so distinct that it might arguably be considered a monotypic subfamily, reliably dating its divergence from

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1012-549: The ability to utilize these pigments for external coloration has evolved independently many times throughout avian evolutionary history. Carotenoid coloration exhibits high levels of sexual dimorphism , with adult male birds generally displaying more vibrant coloration than females of the same species. These differences arise due to the selection of yellow and red coloration in males by female preference . In many species of birds, females invest greater time and resources into raising offspring than their male partners. Therefore, it

1058-669: The bird species Parus major , the more colorfully ornamented males produce sperm that is better protected against oxidative stress due to increased presence of carotenoid antioxidants . However, there is also evidence that attractive male coloration may be a faulty signal of male quality. Among stickleback fish, males that are more attractive to females due to carotenoid colorants appear to under-allocate carotenoids to their germline cells. Since carotinoids are beneficial antioxidants, their under-allocation to germline cells can lead to increased oxidative DNA damage to these cells. Therefore, female sticklebacks may risk fertility and

1104-629: The compounds to be deeply colored yellow, orange, or red. Carotenoids are the dominant pigment in autumn leaf coloration of about 15-30% of tree species, but many plant colors, especially reds and purples, are due to polyphenols . Carotenoids serve two key roles in plants and algae: they absorb light energy for use in photosynthesis , and they provide photoprotection via non-photochemical quenching . Carotenoids that contain unsubstituted beta-ionone rings (including β-carotene , α-carotene , β-cryptoxanthin , and γ-carotene ) have vitamin A activity (meaning that they can be converted to retinol ). In

1150-555: The cytosolic mevalonic acid pathway (MVA) and the plastidic methylerythritol 4-phosphate (MEP). In animals, the production of cholesterol starts by creating IPP and DMAPP using the MVA. For carotenoid production plants use MEP to generate IPP and DMAPP. The MEP pathway results in a 5:1 mixture of IPP:DMAPP. IPP and DMAPP undergo several reactions, resulting in the major carotenoid precursor, geranylgeranyl diphosphate (GGPP). GGPP can be converted into carotenes or xanthophylls by undergoing

1196-468: The development of monoclonal antibodies to trans- lycopene it was possible to localise this carotenoid in different animal and human cells. Beta-carotene , found in pumpkins , sweet potato , carrots and winter squash , is responsible for their orange-yellow colors. Dried carrots have the highest amount of carotene of any food per 100-gram serving, measured in retinol activity equivalents (provitamin A equivalents). Vietnamese gac fruit contains

1242-432: The disappearance of chlorophyll. Carotenoids are responsible for the brilliant yellows and oranges that tint deciduous foliage (such as dying autumn leaves ) of certain hardwood species as hickories , ash , maple , yellow poplar , aspen , birch , black cherry , sycamore , cottonwood , sassafras , and alder . Carotenoids are the dominant pigment in autumn leaf coloration of about 15-30% of tree species. However,

1288-482: The eye, lutein , meso -zeaxanthin , and zeaxanthin are present as macular pigments whose importance in visual function, as of 2016, remains under clinical research . Carotenoids are produced by all photosynthetic organisms and are primarily used as accessory pigments to chlorophyll in the light-harvesting part of photosynthesis. They are highly unsaturated with conjugated double bonds , which enables carotenoids to absorb light of various wavelengths . At

1334-421: The fatty tissues of animals, and exclusively carnivorous animals obtain the compounds from animal fat. In the human diet, absorption of carotenoids is improved when consumed with fat in a meal. Cooking carotenoid-containing vegetables in oil and shredding the vegetable both increase carotenoid bioavailability . The most common carotenoids include lycopene and the vitamin A precursor β-carotene. In plants,

1380-672: The ground. It lays three or four eggs on a ground scrape. The African wattled lapwing is one of the species to which the Agreement on the Conservation of African-Eurasian Migratory Waterbirds ( AEWA ) applies. Lapwing Erythrogonys Vanellus Hoploxypterus and see text Lapwings ( subfamily Vanellinae) are any of various ground-nesting birds ( family Charadriidae ) akin to plovers and dotterels . They range from 10 to 16 inches (25 to 41 cm) in length, and are noted for their slow, irregular wingbeats in flight and

1426-446: The highest known concentration of the carotenoid lycopene . Although green, kale , spinach , collard greens , and turnip greens contain substantial amounts of beta-carotene. The diet of flamingos is rich in carotenoids, imparting the orange-colored feathers of these birds. Reviews of preliminary research in 2015 indicated that foods high in carotenoids may reduce the risk of head and neck cancers and prostate cancer . There

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1472-431: The living Vanellinae or just immediately outside it thus means that their last common ancestor – or even the last common ancestor of plovers and lapwings – almost certainly was a plover-sized bird with a black crown and breast-band, a white feather patch at the wrist, no hallux, and a lipochromic (probably red) bill with a black tip. Its legs were most likely black or the color of the bill's base. The fossil record of

1518-413: The missing hallux (hind toe) are like those of lapwings: it is still not entirely clear whether it is better considered the most basal plover or lapwing. The IOC also recognizes a monotypic genus Hoploxypterus for the pied plover . Many coloration details of the red-kneed dotterel also occur here and there among the living members of the main lapwing clade . Its position as the most basal of

1564-571: The most balanced (lowest energy) state for the carotenoid. Carotenoids defend plants against singlet oxygen , by both energy transfer and by chemical reactions. They also protect plants by quenching triplet chlorophyll. By protecting lipids from free-radical damage, which generate charged lipid peroxides and other oxidised derivatives, carotenoids support crystalline architecture and hydrophobicity of lipoproteins and cellular lipid structures, hence oxygen solubility and its diffusion therein. Like some fatty acids , carotenoids are lipophilic due to

1610-424: The multiple conjugated double bonds determines their color and photophysics. After absorbing a photon, the carotenoid transfers its excited electron to chlorophyll for use in photosynthesis. Upon absorption of light, carotenoids transfer excitation energy to and from chlorophyll . The singlet-singlet energy transfer is a lower energy state transfer and is used during photosynthesis. The triplet-triplet transfer

1656-406: The presence of CTP using the enzyme MEP cytidylyltransferase. CDP-ME is then converted, in the presence of ATP , to 2-phospho-4-(cytidine 5’-diphospho)-2-C-methyl-D-erythritol (CDP-ME2P). The conversion to CDP-ME2P is catalyzed by CDP-ME kinase . Next, CDP-ME2P is converted to 2-C-methyl-D-erythritol 2,4-cyclodiphosphate (MECDP). This reaction occurs when MECDP synthase catalyzes the reaction and CMP

1702-399: The presence of long unsaturated aliphatic chains. As a consequence, carotenoids are typically present in plasma lipoproteins and cellular lipid structures. Carotenoids are located primarily outside the cell nucleus in different cytoplasm organelles, lipid droplets , cytosomes and granules. They have been visualised and quantified by raman spectroscopy in an algal cell. With

1748-524: The reds, the purples, and their blended combinations that decorate autumn foliage usually come from another group of pigments in the cells called anthocyanins . Unlike the carotenoids, these pigments are not present in the leaf throughout the growing season, but are actively produced towards the end of summer. Dietary carotenoids and their metabolic derivatives are responsible for bright yellow to red coloration in birds. Studies estimate that around 2956 modern bird species display carotenoid coloration and that

1794-417: The resulting seedlings. Also, J-protein (J20) and heat shock protein 70 (Hsp70) chaperones are thought to be involved in post-transcriptional regulation of DXS activity, such that mutants with defective J20 activity exhibit reduced DXS enzyme activity while accumulating inactive DXS protein. Regulation may also be caused by external toxins that affect enzymes and proteins required for synthesis. Ketoclomazone

1840-544: The same time, and with the presence of more basal Charadriidae a few million years earlier. However, the assignment of fragmentary fossils to Charadriinae or Vanellinae is not easy. Thus, it is very likely that the charadriid waders originate around the Eocene -Oligocene boundary – roughly 40–30 mya – but nothing more can be said at present. If the Belgian fossil is not a true lapwing, there are actually no Vanellinae fossils known before

1886-759: The same time, the terminal groups regulate the polarity and properties within lipid membranes . Most carotenoids are tetraterpenoids , regular C 40 {\displaystyle {\ce {C40}}} isoprenoids . Several modifications to these structures exist: including cyclization , varying degrees of saturation or unsaturation, and other functional groups . Carotenes typically contain only carbon and hydrogen, i.e., they are hydrocarbons . Prominent members include α-carotene , β-carotene , and lycopene , are known as carotenes . Carotenoids containing oxygen include lutein and zeaxanthin . They are known as xanthophylls . Their color, ranging from pale yellow through bright orange to deep red,

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1932-545: The subfamily remain confused. Some workers have gone so far as to group all the "true" lapwings (except the red-kneed dotterel ) into the single genus Vanellus . Current consensus favors a more moderate position, but it is unclear which genera to split. The Handbook of Birds of the World provisionally places all Vanellinae in Vanellus except the red-kneed dotterel, which is in the monotypic Erythrogonys . Its plesiomorphic habitus resembles that of plovers , but details like

1978-409: The sweet floral smells present in black tea , aged tobacco , grape , and many fruits are due to the aromatic compounds resulting from carotenoid breakdown. Some carotenoids are produced by bacteria to protect themselves from oxidative immune attack. The aureus (golden) pigment that gives some strains of Staphylococcus aureus their name is a carotenoid called staphyloxanthin . This carotenoid

2024-453: The viability of their offspring by choosing redder, but more deteriorated partners with reduced sperm quality. Products of carotenoid degradation such as ionones , damascones and damascenones are also important fragrance chemicals that are used extensively in the perfumes and fragrance industry. Both β-damascenone and β-ionone although low in concentration in rose distillates are the key odor-contributing compounds in flowers. In fact,

2070-488: The xanthophyll lutein is the most abundant carotenoid and its role in preventing age-related eye disease is currently under investigation. Lutein and the other carotenoid pigments found in mature leaves are often not obvious because of the masking presence of chlorophyll . When chlorophyll is not present, as in autumn foliage, the yellows and oranges of the carotenoids are predominant. For the same reason, carotenoid colors often predominate in ripe fruit after being unmasked by

2116-403: The yellow morph of common wall lizards are due to carotenoids. It has been proposed that carotenoids are used in ornamental traits (for extreme examples see puffin birds) because, given their physiological and chemical properties, they can be used as visible indicators of individual health, and hence are used by animals when selecting potential mates. Carotenoids from the diet are stored in

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