Developmental basis for telencephalon expansion in waterfowl: enlargement prior to neurogenesis

Some altricial and some precocial species of birds have evolved enlarged telencephalons compared with other birds. Previous work has shown that finches and parakeets, two species that hatch in an immature (i.e. altricial) state, enlarged their telencephalon by delaying telencephalic neurogenesis. To determine whether species that hatch in a relatively mature (i.e. precocial) state also enlarged their telencephalon by delaying telencephalic neurogenesis, we examined brain development in geese, ducks, turkeys and chickens, which are all precocial. Whereas the telencephalon occupies less than 55 per cent of the brain in chickens and turkeys, it occupies more than 65 per cent in ducks and geese. To determine how these species differences in adult brain region proportions arise during development, we examined brain maturation (i.e. neurogenesis timing) and estimated telencephalon, tectum and medulla volumes from serial Nissl-stained sections in the four species. We found that incubation time predicts the timing of neurogenesis in all major brain regions and that the telencephalon is proportionally larger in ducks and geese before telencephalic neurogenesis begins. These findings demonstrate that the expansion of the telencephalon in ducks and geese is achieved by altering development prior to neurogenesis onset. Thus, precocial and altricial species evolved different developmental strategies to expand their telencephalon.     

Embryonic Calls as Care-soliciting Signals in Budgerigars,Melopsittacus undulatus

Embryonic calls occur 1–3 d before hatching among precocial and some altricial birds. In precocial species, calls may synchronize hatching among siblings or, in semi-precocial species, elicit parental attention to, and often thermoregulation of, the hatching egg. Much less is known about the functional significance of calls in fully altricial species. In this study, naturalistic observations and laboratory experiments were used to document factors affecting calling and the parental responses to calls in one altricial species, the budgerigar Melopsittacus undulatus. Budgerigar chicks hatch asynchronously and vocalize 24–48 h before hatching. Embryonic calling rates increase at higher egg temperatures, and also as embryos near hatching. Parents easily locate a calling egg in their clutch, even among a large brood of much older, vocalizing nestlings. Furthermore, they actively assist in the last stages of hatching by helping to break the shell along the crack in the egg. Both observational and experimental evidence suggests that embryonic vocalizations are distinctive signals that increase parental attention and care, and may stimulate hatching assistance to a calling egg.     

Establishment of the fatty acid profile of the brain of the king penguin (Aptenodytes patagonicus) at hatch: effects of a yolk that is naturally rich in n-3 polyunsaturates

Because the yolk lipids of the king penguin (Aptenodytes patagonicus) contain the highest concentrations of long-chain n-3 polyunsaturated fatty acids yet reported for an avian species, the consequences for the establishment of the brain's fatty acid profile in the embryo were investigated. To place the results in context, the fatty acid compositions of yolk lipid and brain phospholipid of the king penguin were compared with those from three other species of free-living birds. The proportions of docosahexaenoic acid (22:6n-3; DHA) in the total lipid of the initial yolks for the Canada goose (Branta canadensis), mallard (Anas platyrhynchos), moorhen (Gallinula chloropus), and king penguin were (% w/w of fatty acids) 1.0+/-0.1, 1.9+/-0.2, 3.3+/-0.1, and 5.9+/-0.2, respectively. The respective concentrations of DHA (% w/w of phospholipid fatty acids) in brains of the newly hatched chicks of these same species were 18.5+/-0.2, 19.6+/-0.7, 16.9+/-0.4, and 17.6+/-0.1. Thus, the natural interspecies diversity in yolk fatty acid profiles does not necessarily produce major differences in the DHA content of the developing brain. Only about 1% of the amount of DHA initially present in the yolk was recovered in the brain of the penguin at hatch. There was no preferential uptake of DHA from the yolk during development of the king penguin.     

Altricial and precocial developmental features in modern meat-type chicks

Variability of egg weight, egg yolk content, neonatal growth rate and relationships of these parameters were studied in meat-type chicks. As it had been established the level of variability in neonatal growth traits was greater than variability of the egg morphology parameters. Egg weight had stronger influence on the chicks' neonatal growth rate than egg yolk content did. Low egg size was associated with limited neonatal growth rate variability, declined chick weight at hatching and increased relative growth rate throughout four days post hatch. Comparison of egg morphological parameters in two species having the same female definitive body weight--meat-type domestic fowl (precocial type) and brown pelican (altricial type) has shown, that, in contrary to predicted on the basis of avian developmental typology, egg weight to female body ratio was greater in brown pelican, egg yolk content was equal in both species.     

Brain size, development and metabolism in birds and mammals

Recent hypotheses that variation in brain size among birds and mammals result from differences in metabolic allocation during ontogeny are tested.
Indices of embryonic and post-embryonic brain growth are defined. Precocial birds and mammals have high embryonic brain growth indices which are compensated for by low post-embryonic indices (with the exception of Homo supiens ). In contrast, altricial birds and mammals have low embryonic and high post-embryonic indices. Altricial birds have relatively small brains at hatching and develop relatively large brains as adults, but among mammals there is no equivalent correlation between variation in adult relative brain sizes and state of neonatal development.
Compensatory brain development in both birds and mammals is associated with compensatory parental metabolic allocation. In comparison with altricial development, precocial development is characterized by higher levels of brain growth and parental metabolic allocation prior to hatching or birth and lower levels subsequently. Differences between degrees of postnatal investment by the parents in the young of precocial birds versus precocial mammals may result in the different patterns of adult brain size associated with precociality versus altriciality in the two groups.
The allometric exponent scaling brain on body size differs among taxonomic levels in birds. The exponent is higher for some parts of the brain than others, irrespective of taxonomic level. Unlike mammals, the exponents for birds do not show a general increase with taxonomic level. These pattcrns call into question recent interpretations of the allometric exponent in birds. and the reason for changes in exponent with taxonomic level.     

Changes in catecholamines and dopaminergic metabolites in pigeon brain during development from the late embryonic stage toward hatch

While brain development during embryogenesis has been extensively studied in precocial birds, there is no information available on altricial birds. Thus, the concentrations of the catecholamines norepinephrine (NE), epinephrine (E), and dopamine (DA), and the dopaminergic metabolites 3,4-dihydroxyphenylacetic acid (DOPAC) and 4-hydroxy-3-methoxyphenylacetic acid (HVA) were determined at several stages during the late embryonic period (E13, E14, E15, E16, E17 and E18) and the day-of-hatch (P0) in the pigeon telencephalon, cerebellum, optic lobe, and brainstem. The concentrations of all catecholamines were higher than those reported in chicken embryos. During embryogenesis, NE, E, DOPAC and HVA concentrations in the various brain parts increased throughout embryonic development until shortly before hatching at which time they decreased. DA, however, continued to increase through hatching in the brainstem, and the changes in DA concentrations varied in several brain parts. In conclusion, catecholamine concentrations in the various brain parts tended to increase with embryonic age, and the concentrations were higher than those in chickens. Furthermore, brain catcholamine metabolism changed at hatch in pigeons.     

A comparative study of embryonic development of some bird species with different patterns of postnatal growth

Some studies show that birds with high postnatal growth rates (e.g. altricial species) are characterized by a rapid early development of "supply" organs, such as digestive organs. Birds with low postnatal growth rates (e.g. precocial species) exhibit a slower early development of these organs and a more rapid early development of other "demand" organs, such as brain, muscles, skeleton and feathers. To test whether these differences can be traced back to early embryonic development and whether they can be associated with changes in developmental timing, i.e. heterochrony, we compared embryos of the precocial quail and the altricial fieldfare, two bird species with low and high postnatal growth rates, respectively. We used classical staging techniques that use developmental landmarks to categorize embryonic maturity as well as morphological measurements. These techniques were combined with immune detection of muscle specific proteins in the somites. Our data showed that the anlagen of the head, brain and eyes develop earlier in the quail than in the fieldfare in contrast to the gut which develops earlier in the fieldfare than in the quail. Our data also showed that the quail and the fieldfare displayed different rates of myotome formation in the somites which contribute to muscle formation in the limbs and thorax. We believe these observations are connected with important differences in neonatal characteristics, such as the size of the brain, eyes, organs for locomotion and digestion. This leads us to the conclusion that selection for late ontogenetic characteristics can alter early embryonic development and that growth rate is of fundamental importance for the patterning of avian embryonic development. It also appears that this comparative system offers excellent opportunities to test hypotheses about heterochrony.     

Effects of egg size on Double-crested Cormorant (Phalacrocorax auritus) egg composition and hatchling phenotype

Maternal investment of yolk and albumen in avian eggs varies with egg mass and contributes to variation in hatchling mass. Here we use the natural variation in mass and composition of Double-crested Cormorant (Phalacrocorax auritus) eggs to examine consequences of variation in yolk and albumen mass on hatchling phenotype. The Double-crested Cormorant, a large bird with altricial young, lays eggs ranging in mass from 40 to 60 g and containing an average of 82% albumen and 18% yolk. Variation in Cormorant egg mass arises primarily from variation in the amount of albumen and water in the eggs; yolk mass remains relatively constant, contributing only 10% to egg mass variation. Likewise, variation in hatchling mass correlates positively with albumen mass and albumen solids contribute to hatchling dry mass. Thus, variation in Cormorant egg mass is primarily the result of variation in the amount of egg albumen, which contributes most to variation in hatchling mass. Similarities in egg composition of altricial birds, along with data presented here, suggest that variation in hatchling mass of all altricial birds may depend most on the amount of egg albumen, unlike species with precocial young that hatch from eggs with substantially more yolk.     

Eggshell structure, mode of development and growth rate in birds

The shell of an egg contributes to successful embryogenesis in many ways, such as through protection, respiration and water exchange. The shell is also the major source of calcium for the development of high-calcium consuming organs, e.g. the skeleton, muscles and brain. Some studies show, moreover, that growth rate may play a fundamental role in the pattern of skeletal development in birds: the faster the growth the less ossified the skeleton is at hatching. We predicted, therefore, that slow (precocial) and fast (altricial) growing bird species should lay eggs encased in shells with different structures adapted to support different rates of calcium removal by developing embryos. We tested this prediction by comparing the fine structure of the inner eggshell surface (mammillary layer) from 36 bird species belonging to 18 orders ranging from Struthioniformes to Passeriformes. Using scanning electron microscopy, we compared the mammillary layer of both non-incubated eggs and eggs at the time of hatching, i.e., before and after embryonic development and the accompanying calcium removal. The results were consistent with the prediction, i.e., the number of mammillary tips per unit of surface area was associated with mode of development and growth rate. The number was higher, and calcium removal was also more extensive, in shells from precocial bird species than in shells from altricial bird species.     

Ontogenetic patterns of constitutive immune parameters in altricial house sparrows

Innate immune functions are proposed to develop rapidly post‐hatch in altricial nestlings, compared with adaptive immune defenses that require development of receptor specificity and memory. Studies of ontogenetic changes in altricial birds have been few until relatively recently and often do not encompass the entire developmental period. We examined the patterns of development in constitutive innate and adaptive immune indices in house sparrow nestlings (3, 6, 9 and 12 days (d) post‐hatch), hatch‐year birds and adults. Lysozyme activity significantly decreased with age, likely representing catabolism of maternal investment of lysozyme in the egg albumen. Levels of total IgY (indexing adaptive immune function), as well as agglutination and lysis (indexing innate immune function), increased throughout the nestling period, but were significantly below levels found in fully‐grown birds at the time of fledging. There were no significant differences between hatch‐year birds and adults in these measures, indicating that rapid, full maturation occurs early in the post‐fledging period. In combination with previous studies, these data highlight the importance of sampling fledglings to assess full immune ontogeny and suggest that fledgling birds may be more vulnerable to infection than adults.     

Strategies of mass change in breeding birds

Birds experience strikingly different patterns of mass change during breeding. In some species with uniparental incubation, the incubating parents (mostly females) lose mass and attain their lowest point when the chicks hatch (Incubatory mass loss strategy = IML). In species with shared incubation, or with intense incubation feeding, the incubating parents maintain or increase in body mass without reducing attentiveness levels (Incubatory mass constancy = IMC). In other species, uniparental incubation with IMC is associated with reduced levels of attentiveness. The mobilization of fat stores or its preservation are involved in the two strategies. IML leads to mass increases after hatching of the young, while the opposite is true for IMC. The non-incubating sex in uniparental species does not experience significant mass changes during breeding. Fasting endurance, predation risk and mode of development are proposed as the main selective factors determining strategy. Latitude, climate, diet and food availability interact with the main factors. From a revision of the literature we can deduce that IML is mainly found among large birds with precocial development, while IMC is typical of smaller species with altricial development. Proportional mass losses are positively correlated with body mass in IML-species, as well as in precocial species, where body mass explains more than 70% of the variation in proportional mass change. Incubation periods increase with body mass in uniparental IMC-species, but not in IML-species. IML is thus associated to fast embryonic growth in large species. Successful raising of highly-dependent young in altricial and semialtricial species apparently depends on one of the parents retaining reserves until hatching. Subsequent mass losses may be necessary to maintain the brooding parent through a period when nestlings require heat, insulation and food. Patterns of mass change are not mere consequences of reproductive stress but the outcome of adaptive compromises between different selective factors and constitute an important aspect of the breeding biology and life history of birds.     

Patterns of mobilization of calcium,magnesium, and phosphorus by embryonic yellow-headed blackbirds(Xanthocephalus xanthocephalus)

Summary Embryonic blackbirds(Xanthocephalus xanthocephalus) obtain most of their calcium from the eggshell (85 90%), but all of their phosphorus comes from reserves in the yolk (80–85%) and albumen (15–20%). Approximately equal amounts of magnesium are supplied by the eggshell, the yolk, and the albumen. Yolk is depleted of magnesium and phosphorus during embryogenesis, but excess calcium absorbed from the eggeshell is stored in the yolk. Consequently reserves of calcium in the yolk actually increase 8-fold during embryonic development. Our results reveal that altricial birds manifest patterns of mobilization and deposition of calcium and other elements similar to those described for precocial species. Evolution of altriciality from precocity evidently did not entail major changes in how embryonic birds meet the challenge of obtaining the calcium, magnesium, and phosphorus required for development.     

Metabolic fates of yolk lipid and individual fatty acids during embryonic development of the coot and moorhen

There is currently little information regarding the metabolic fates of yolk lipid and individual fatty acids during embryonic development of free-living avian species. Here we report the pattern of lipid utilization during embryonic development of the coot (Fulica atra) and the moorhen (Gallinula chloropus), two related species producing precocial offspring from eggs with a distinctive fatty acid composition and with an incubation period similar to that of the chicken. By the time of hatching, the proportions of the initial yolk lipid that had been transferred to the embryo were 88.2% and 79.8% for the coot and moorhen respectively. During the whole incubation period, 42.9% and 40.0% of the initial yolk lipid of the coot and moorhen respectively were lost from the system due to oxidation for energy, equating to 47.8% and 50.0% respectively of the actual amount of lipid transferred over this time. Thus, the lipid received by the embryos of both species is partitioned almost equally between the alternative fates of energy metabolism and incorporation into tissue lipids. In the coot, this 50:50 split between oxidation and tissue formation was maintained during the hatching process. The proportions of arachidonic (20:4n-6) and docosahexaenoic (22:6n-3) in the yolk lipids of these species were 2.5-3.5 times higher than in eggs of domestic poultry. In contrast to the situation in the chicken, there was no preferential uptake of 22:6n-3 from the yolk during coot and moorhen development. The fatty acid compositions of the whole body lipids of the coot and moorhen hatchlings were almost identical to those of the initial yolks indicating that, unlike the chicken, these species display relatively little overall biomagnification of 20:4n-6 and 22:6n-6 during development. It is suggested that the yolk fatty acid profiles of the coot and moorhen are particularly well matched to the requirements of the embryo, reducing the need for selective uptake of 22:6n-3 and for the overall biomagnification of 22:6n-3 and 20:4n-6.     

Comparative growth rates and oxygen consumption in young Galliformes

The high correlation between growth rate and adult body weight has been much more thoroughly documented for altricial birds than for precocial species. This paper gathers data from the literature for precocial Galliformes and also reports new growth data on six galliform species for analysis. The onset of homeothermic ability is investigated in Galliformes over a range of body size. The results confirm that (1) large species' chicks grow at a slower rate than those of smaller species, and (2) larger species' chicks can thermoregulate earlier than smaller species' chicks under cold stress situations. Published embryonic body weights are also analysed to determine when growth rate differences appear in the development of precocial species. No interspecific differences appeared in the relative growth rates of embryos, and therefore species body size does not appear to influence growth rate before hatching.     

Differential incorporation of docosahexaenoic and arachidonic acids by the yolk sac membrane of the avian embryo

During avian development, lipoproteins derived from yolk lipid are assembled in the yolk sac membrane (YSM) for secretion into the embryonic circulation. To investigate how yolk polyunsaturated fatty acids, essential for the development of certain tissues, are distributed among the lipid classes of the lipoproteins, pieces of YSM were incubated in vitro with [14C]arachidonic and [14C]docosahexaenoic acids (DHA). There was a marked difference in the partitioning of these two precursors among the lipid classes of the tissue. Of the radioactivity incorporated into total lipid from [14C]-arachidonic acid during 1 h of incubation, 67.3% was esterified as phospholipid and 29.5% as triacylglycerol. In contrast, only 14.6% of the label incorporated from [14C]-DHA was esterified as phospholipid, whereas 73.2% was recovered in triacylglycerol. This pattern of differential partitioning was observed at all time points and across a 20-fold range of fatty acid concentrations. There was no evidence for conversion of the radioactive arachidonic and DHAs to other fatty acids prior to incorporation into tissue lipids. It is suggested that the selective incorporation of yolk-derived DHA into the triacylglycerol of secreted lipoproteins represents part of a mechanism for directing this polyunsaturate to particular embryonic tissues.     

The relationship between the fatty acid profiles of the yolk and the embryonic tissue lipids: A comparison between the lesser black backed gull (Larus fuscus) and the pheasant (Phasianus colchicus)

Although substantial information is available regarding the fatty acid composition of lipids of the yolk and of the developing tissues of the chicken embryo, there is little knowledge on this topic for other avian species. The aim of the present study was to compare the yolk and embryonic tissue fatty acid profiles for a species selecting its food in the wild (the lesser black backed gull) with one fed on a standard commercial diet (the commercially reared pheasant). The fatty acid compositions of the yolk lipids were determined, and major differences were observed between the two species. In particular, the phospholipid of the gull yolk was enriched in 20:4n-6 and 22:6n-3 (18.8 and 7.1%, respectively, by weight of total fatty acids) in comparison with the pheasant (4.0 and 4.1%, respectively). The fatty acid compositions of the embryonic tissues were determined using eggs incubated in the laboratory. For the liver and heart, the fatty acid composition of the lipids in the two species reflected the initial yolk composition, with the gull tissue lipids generally containing higher proportions of 20:4n-6 and 22:6n-3 than those of the pheasant. In contrast, the fatty acid profiles of the brain phospholipid were essentially identical in the two species, with 20:4n-6 and 22:6n-3 comprising approximately 9 and 17%, respectively, of total fatty acids in both cases.     

Composition, accumulation and utilization of yolk lipids in teleost fish

Lipid reserves in teleost eggs are stored in lipoprotein yolk and, in some species, a discrete oil globule. Lipoprotein yolk lipids are primarily polar lipids, especially phosphatidylcholine and phosphatidylethanolamine (PE), and are rich in (n–3) polyunsaturated fatty acids, especially 22:6(n–3) (docosahexaenoic acid, DHA). Oil consists of neutral lipids and is rich in monounsaturated fatty acids (MUFA). Egg lipids are derived from dietary fatty acid, fatty acid mobilized from reserves and possibly fatty acid synthesized de novo. There is selective incorporation of essential fatty acids, particularly DHA, into yolk lipids and discrimination against incorporation of 22:1(n–11). Lipid is delivered to the oocyte by vitellogenin, which is rich in polar lipids, and likely also by other lipoproteins, especially very low density lipoprotein, which is rich in triacylglycerol (TAG). All classes of lipid may be used as fuel during embryonic and larval development and MUFA are preferred fatty acids for catabolism by embryos. Catabolism of oil globules is frequently delayed until latter stages of development. In some species, DHA derived from hydrolysis of phospholipid may be conserved by transfer to the neutral lipid. Recent work has expanded knowledge of the role of DHA in membrane structure, especially in neural tissue, and molecular species analysis has indicated that PE containing sn-1 oleic acid is a prime contributor to membrane fluidity. The results of this type of study provide an explanation for the selection pressures that influence yolk lipid composition. Future work ought to expand knowledge of specific roles of individual fatty acids in embryos along with knowledge of the ecological physiology of ovarian recrudescence, environmental influences on vitellogenin and yolk lipid composition, and the control of yolk lipid accumulation and utilization.     

Ontogenetic development of the ophthalmic rete in relation to brain cooling in chickens and pigeons

The brain cooling capacity of the altricial pigeon increases during posthatching growth at a higher rate than that of the precocial duck and chicken. To determine if this difference between the altricial and the precocial modes of development can be related to growth rates of the vascular heat exchanger involved in brain cooling (the ophthalmic rete), we performed a morphometric analysis of this structure during the post-hatching maturation of the three species. The number of vascular units in the rete did not change during development but differed significantly among species. The retia continued to grow in length and diameter in an exponential relation with body mass at similar rates in all species. The surface area of the retial arteries, which reflects the area available for countercurrent heat exchange, also increased exponentially with body mass, but without significant differences among the three species. However, the effectiveness of the rete in brain cooling, as indicated by the degree of brain cooling per unit of heat-exchange area in the rete, was higher in the altricial pigeon than in the precocial chicken and duck. It is concluded that the posthatching morphometric changes in the ophthalmic rete (rete ophthalmicum) are important for the development of brain cooling capacity, but cannot solely explain differences in brain cooling between growing altricial and precocial birds. These differences are most likely related to differences in the maturation of the central thermoregulatory control system and the peripheral effector mechanisms among the two groups of birds.     

Distribution of lipids from the yolk to the tissues during development of the water python (<Emphasis Type="Italic">Liasis fuscus</Emphasis>)

Energy metabolism during embryonic development of snakes differs in several respects from the patterns displayed by other reptiles. There are, however, no previous reports describing the main energy source for development, the yolk lipids, in snake eggs. There is also no information on the distribution of yolk fatty acids to the tissues during snake development. In eggs of the water python (Liasis fuscus), we report that triacylglycerol, phospholipid, cholesteryl ester and free cholesterol, respectively, form 70.3%, 14.1%, 5.7% and 2.1% of the total lipid. The main polyunsaturate of the yolk lipid classes is 18:2n-6. The yolk phospholipid contains 20:4n-6 and 22:6n-3 at 13.0% and 3.6% (w/w), respectively. Approximately 10% and 30% of the initial egg lipids are respectively recovered in the residual yolk and the fat body of the hatchling. A major function of yolk lipid is, therefore, to provision the neonate with large energy reserves. The proportion of 22:6n-3 in brain phospholipid of the hatchling is 11.1% (w/w): this represents only 0.24% of the amount of 22:6n-3 originally present in the egg. This also contrasts with values for free-living avian species where the proportion of DHA in neonatal brain phospholipid is 16–19%. In the liver of the newly hatched python, triacylglycerol, phospholipid and cholesteryl ester, respectively, form 68.2%, 7.7% and 14.3% of total lipid. This contrasts with embryos of birds where cholesteryl ester forms up to 80% of total liver lipid and suggests that the mechanism of lipid transfer in the water python embryo differs in some respects from the avian situation.Abbreviations ARA arachidonic acid - DHA docosahexaenoic acidCommunicated by G. Heldmaier     

Development of parameters influencing blood oxygen carrying capacity in the welcome swallow and fairy martin

Welcome swallow (Hirundo neoxena) and fairy martin (Petrochelidon ariel) nestlings develop relatively slowly. They exceed adult mass early in development, then lose weight and fledge at about adult mass, i.e. have a mass-overshoot-recession (Mor) growth profile. Development of parameters influencing blood oxygen carrying capacity (O2Cap) was documented to determine if O2Cap also reached a plateau substantially before fledging or increased continuously throughout nestling development. Hematocrit (Hct), erythrocyte count (RBC) and whole blood haemoglobin (Hb) increased 1.8- to 2.8-fold, so that O2Cap doubled during development in both species. Increase in Hct, Hb and RBC was not continuous, peak values occurring well before fledging, in contrast to passerines with standard growth profiles in which the increase occurs throughout nestling development and peak values occur at fledging. However, the timing of O2Cap increase differed from that in some other Mor species (e.g. shearwaters). Mean erythrocyte volume (MCV) decreased linearly throughout development by 30-41%, but mean erythrocyte haemoglobin content (MCH) remained constant, so that mean erythrocyte haemoglobin concentration (MCHC) increased linearly 1.3- to 1.5-fold. Possible reasons for the apparent differences in the timing of O2Cap increase between rapidly and slowly growing altricial species and among MOR species are discussed.