13.3: Evolution of Vertebrata/Craniata

Learning Objectives

By the end of this section, you will be able to do the following:

• Identify the derived characters of craniates that sets them apart from other chordates
• Describe the developmental fate of the notochord in vertebrates

 

The vertebrates exhibit two major innovations in their evolution from the invertebrate chordates. These innovations may be associated with the whole genome duplications that resulted in a quadruplication of the basic chordate genome, including the Hox gene loci that regulate the placement of structures along the three axes of the body. One of the first major steps was the emergence of the quadrupeds in the form of the amphibians. A second step was the evolution of the amniotic egg, which, similar to the evolution of pollen and seeds in plants, freed terrestrial animals from their dependence on water for fertilization and embryonic development. Within the amniotes, modifications of keratinous epidermal structures have given rise to scales, claws, hair, and feathers. The scales of reptiles sealed their skins against water loss, while hair and feathers provided insulation to support the evolution of endothermy, as well as served other functions such as camouflage and mate attraction in the vertebrate lineages that led to birds and mammals.

Currently, a number of vertebrate species face extinction primarily due to habitat loss and pollution. According to the International Union for the Conservation of Nature, more than 6,000 vertebrate species are classified as threatened. Amphibians and mammals are the classes with the greatest percentage of threatened species, with 29 percent of all amphibians and 21 percent of all mammals classified as threatened. Attempts are being made around the world to prevent the extinction of threatened species. For example, the Biodiversity Action Plan is an international program, ratified by 188 countries, which is designed to protect species and habitats.

 

---

Characteristics of Vertebrata (Craniata)

Members of the phylum Craniata/Vertebrata display the five characteristic features of the chordates; however, members of this group also share derived characteristics that distinguish them from invertebrate chordates. Vertebrates are named for the vertebral column, composed of vertebrae—a series of separate, irregularly shaped bones joined together to form a backbone (Figure 29.8). Initially, the vertebrae form in segments around the embryonic notochord, but eventually replace it in adults. In most derived vertebrates, the notochord becomes the nucleus pulposus of the intervertebral discs that cushion and support adjacent vertebrae. 

Figure 29.8 A vertebrate skeleton. Vertebrata are characterized by the presence of a backbone, such as the one that runs through the middle of this fish. All vertebrates are in the Craniata clade and have a cranium. (credit: Ernest V. More; taken at Smithsonian Museum of Natural History, Washington, D.C.)

 

We should note that not all organisms called "vertebrates" technically have vertebrae. The hagfish (Myxini) lack a backbone. Therefore, Craniata may be a more accurate name for the group than Vertebrata, because all organisms in this group do have a cranium. In addition to a cranium, organisms within Vertebrata/Craniata also possess several other defining characteristics: hemoglobin, a liver, kidneys, and a well-developed brain.

 

Cranium

A cranium is a bony, cartilaginous, or fibrous structure surrounding the brain, jaw, and facial bones (Figure 29.7). Most bilaterally symmetrical animals have a head; of these, those that have a cranium comprise the clade Craniata/Vertebrata, which includes the primitively jawless Myxini (hagfishes), Petromyzontida (lampreys), and all of the organisms called “vertebrates.” (We should note that )

Figure 29.7 A craniate skull. The subphylum Craniata (or Vertebrata), including this placoderm fish (Dunkleosteus sp.), are characterized by the presence of a cranium, mandible, and other facial bones. (credit: “Steveoc 86”/Wikimedia Commons)

 

Hemoglobin

Hemoglobin, or Hb, is a protein molecule found in red blood cells (erythrocytes) made of four subunits: two alpha subunits and two beta subunits (Figure 39.19). Each subunit surrounds a central heme group that contains iron and binds one oxygen molecule, allowing each hemoglobin molecule to bind four oxygen molecules. Molecules with more oxygen bound to the heme groups are brighter red. As a result, oxygenated arterial blood where the Hb is carrying four oxygen molecules is bright red, while venous blood that is deoxygenated is darker red.

Figure 39.19 The protein inside (a) red blood cells that carries oxygen to cells and carbon dioxide to the lungs is (b) hemoglobin. Hemoglobin is made up of four symmetrical subunits and four heme groups. Iron associated with the heme binds oxygen. It is the iron in hemoglobin that gives blood its red color.

 

Liver

The liver is the largest internal organ in humans and it plays a very important role in digestion of fats and detoxifying blood. The liver produces bile, a digestive juice that is required for the breakdown of fatty components of the food in the duodenum. The liver also processes the vitamins and fats and synthesizes many plasma proteins.

 

Kidneys

The kidneys, illustrated in Figure 41.4, are a pair of bean-shaped structures that are located just below and posterior to the liver in the peritoneal cavity. The adrenal glands sit on top of each kidney and are also called the suprarenal glands. Kidneys filter blood and purify it. All the blood in the human body is filtered many times a day by the kidneys; these organs use up almost 25 percent of the oxygen absorbed through the lungs to perform this function. Oxygen allows the kidney cells to efficiently manufacture chemical energy in the form of ATP through aerobic respiration. The filtrate coming out of the kidneys is called urine.

Figure 41.4 Kidneys filter the blood, producing urine that is stored in the bladder prior to elimination through the urethra. (credit: modification of work by NCI)

 

Kidneys filter blood in a three-step process. The functional unit of the kidney is the nephron, illustrated in Figure 41.6. Each kidney is made up of over one million nephrons. A nephron consists of three parts—a renal corpuscle, a renal tubule, and the associated capillary network.

Visual Connection

Figure 41.6 The nephron is the functional unit of the kidney. The glomerulus and convoluted tubules are located in the kidney cortex, while collecting ducts are located in the pyramids of the medulla. (credit: modification of work by NIDDK)

 

First, the nephrons filter blood that runs through the capillary network in the glomerulus. Almost all solutes, except for proteins, are filtered out into the glomerulus by a process called glomerular filtration. Second, the filtrate is collected in the renal tubules. Most of the solutes get reabsorbed in the PCT by a process called tubular reabsorption. In the loop of Henle, the filtrate continues to exchange solutes and water with the renal medulla and the peritubular capillary network. Water is also reabsorbed during this step. Then, additional solutes and wastes are secreted into the kidney tubules during tubular secretion, which is, in essence, the opposite process to tubular reabsorption. The collecting ducts collect filtrate coming from the nephrons and fuse in the medullary papillae. From here, the papillae deliver the filtrate, now called urine, into the minor calyces that eventually connect to the ureters through the renal pelvis. This entire process is illustrated in Figure 41.7.

Figure 41.7 Each part of the nephron performs a different function in filtering waste and maintaining homeostatic balance. (1) The glomerulus forces small solutes out of the blood by pressure. (2) The proximal convoluted tubule reabsorbs ions, water, and nutrients from the filtrate into the interstitial fluid, and actively transports toxins and drugs from the interstitial fluid into the filtrate. The proximal convoluted tubule also adjusts blood pH by selectively secreting ammonia (NH₃) into the filtrate, where it reacts with H⁺ to form NH₄⁺. The more acidic the filtrate, the more ammonia is secreted. (3) The descending loop of Henle is lined with cells containing aquaporins that allow water to pass from the filtrate into the interstitial fluid. (4) In the thin part of the ascending loop of Henle, Na⁺ and Cl^- ions diffuse into the interstitial fluid. In the thick part, these same ions are actively transported into the interstitial fluid. Because salt but not water is lost, the filtrate becomes more dilute as it travels up the limb. (5) In the distal convoluted tubule, K⁺ and H⁺ ions are selectively secreted into the filtrate, while Na⁺, Cl^-, and HCO₃^- ions are reabsorbed to maintain pH and electrolyte balance in the blood. (6) The collecting duct reabsorbs solutes and water from the filtrate, forming dilute urine. (credit: modification of work by NIDDK)

 

Brain

The brain is the part of the central nervous system that is contained in the cranial cavity of the skull. Brain anatomy and structure varies greatly between vertebrate groups. In mammals and reptiles (including birds), the brain includes the cerebral cortex, limbic system, thalamus, hypothalamus, and cerebellum. 

 

Cerebral Cortex

The outermost part of the brain is a thick piece of nervous system tissue called the cerebral cortex, which is folded into hills called gyri (singular: gyrus) and valleys called sulci (singular: sulcus). The cortex is made up of two hemispheres—right and left—which are separated by a large sulcus. A thick fiber bundle called the corpus callosum (Latin: “tough body”) connects the two hemispheres and allows information to be passed from one side to the other. Although there are some brain functions that are localized more to one hemisphere than the other, the functions of the two hemispheres are largely redundant. In fact, sometimes (very rarely) an entire hemisphere is removed to treat severe epilepsy. 

Each cortical hemisphere contains regions called lobes that are involved in different functions: frontal, parietal, temporal, and occipital. Figure 35.21 illustrates these four lobes of the human cerebral cortex. The frontal lobe is located at the front of the brain, over the eyes. This lobe contains the olfactory bulb, which processes smells. The frontal lobe also contains the motor cortex, which is important for planning and implementing movement. Neurons in the frontal lobe also control cognitive functions like maintaining attention, speech, and decision-making. Studies of humans who have damaged their frontal lobes show that parts of this area are involved in personality, socialization, and assessing risk. The parietal lobe is located at the top of the brain. Neurons in the parietal lobe are involved in speech and also reading. Two of the parietal lobe’s main functions are processing somatosensation—touch sensations like pressure, pain, heat, cold—and processing proprioception—the sense of how parts of the body are oriented in space. The occipital lobe is located at the back of the brain. It is primarily involved in vision—seeing, recognizing, and identifying the visual world. The temporal lobe is located at the base of the brain by your ears and is primarily involved in processing and interpreting sounds. It also contains the hippocampus (Greek for “seahorse”)—a structure that processes memory formation. 

Figure 35.21 The human cerebral cortex includes the frontal, parietal, temporal, and occipital lobes.

 

Evolution Connection

Cerebral Cortex: Compared to other vertebrates, mammals have exceptionally large brains for their body size. An entire alligator’s brain, for example, would fill about one and a half teaspoons. This increase in brain to body size ratio is especially pronounced in apes, whales, and dolphins. While this increase in overall brain size doubtlessly played a role in the evolution of complex behaviors unique to mammals, it does not tell the whole story. Scientists have found a relationship between the relatively high surface area of the cortex and the intelligence and complex social behaviors exhibited by some mammals. This increased surface area is due, in part, to increased folding of the cortical sheet (more sulci and gyri). For example, a rat cortex is very smooth with very few sulci and gyri. Cat and sheep cortices have more sulci and gyri. Chimps, humans, and dolphins have even more.

Figure 35.23 Mammals have larger brain-to-body ratios than other vertebrates. Within mammals, increased cortical folding and surface area is correlated with complex behavior.

 

Thalamus

The thalamus (Greek for “inner chamber”), illustrated in Figure 35.24, acts as a gateway to and from the cortex. It receives sensory and motor inputs from the body and also receives feedback from the cortex. This feedback mechanism can modulate conscious awareness of sensory and motor inputs depending on the attention and arousal state of the animal. The thalamus helps regulate consciousness, arousal, and sleep states.

Figure 35.24 The limbic system regulates emotion and other behaviors. It includes parts of the cerebral cortex located near the center of the brain, including the cingulate gyrus and the hippocampus as well as the thalamus, hypothalamus, and amygdala.

 

Hypothalamus

Below the thalamus is the hypothalamus, shown in Figure 35.24. The hypothalamus controls the endocrine system by sending signals to the pituitary gland, a pea-sized endocrine gland that releases several different hormones that affect other glands as well as other cells. This relationship means that the hypothalamus regulates important behaviors that are controlled by these hormones. The hypothalamus is the body’s thermostat—it makes sure key functions like food and water intake, energy expenditure, and body temperature are kept at appropriate levels. Neurons within the hypothalamus also regulate circadian rhythms, sometimes called sleep cycles.

 

Limbic System

The limbic system is a connected set of structures that regulates emotion, as well as behaviors related to fear and motivation. It plays a role in memory formation and includes parts of the thalamus and hypothalamus as well as the hippocampus. One important structure within the limbic system is a temporal lobe structure called the amygdala (Greek for “almond”), illustrated in Figure 35.24. The two amygdala are important both for the sensation of fear and for recognizing fearful faces. The cingulate gyrus helps regulate emotions and pain.

 

Cerebellum

The cerebellum (Latin for “little brain”), shown in Figure 35.21, sits at the base of the brain on top of the brainstem. The cerebellum controls balance and aids in coordinating movement and learning new motor tasks.

 

Brainstem

The brainstem, illustrated in Figure 35.21, connects the rest of the brain with the spinal cord. It consists of the midbrain, medulla oblongata, and the pons. Motor and sensory neurons extend through the brainstem allowing for the relay of signals between the brain and spinal cord. Ascending neural pathways cross in this section of the brain allowing the left hemisphere of the cerebrum to control the right side of the body and vice versa. The brainstem coordinates motor control signals sent from the brain to the body. The brainstem controls several important functions of the body including alertness, arousal, breathing, blood pressure, digestion, heart rate, swallowing, walking, and sensory and motor information integration.

 

---

Diversity of Vertebrata/Craniata

The relationship of the vertebrates to the invertebrate chordates has been a matter of contention, but although these cladistic relationships are still being examined, it appears that the Craniata/Vertebrata are a monophyletic group that shares the five basic chordate characteristics with the other two subphyla, Urochordata and Cephalochordata. Traditional phylogenies place the cephalochordates as a sister clade to the chordates, a view that has been supported by most current molecular analyses. This hypothesis is further supported by the discovery of a fossil in China from the genus Haikouella. This organism seems to be an intermediate form between cephalochordates and vertebrates. The Haikouella fossils are about 530 million years old and appear similar to modern lancelets. These organisms had a brain and eyes, as do vertebrates, but lack the skull found in craniates.¹ This evidence suggests that vertebrates arose during the Cambrian explosion.

Vertebrates are the largest group of chordates, with more than 62,000 living species, which are grouped based on anatomical and physiological traits. More than one classification and naming scheme is used for these animals. Here we will consider the traditional groups Agnatha, Chondrichthyes, Osteichthyes, Amphibia, Reptilia, Aves, and Mammalia, which constitute classes in the subphylum Vertebrata/Craniata. Virtually all modern cladists classify birds within Reptilia, which correctly reflects their evolutionary heritage. Thus, we now have the nonavian reptiles and the avian reptiles in our reptilian classification. We consider them separately only for convenience. Further, we will consider hagfishes and lampreys together as jawless fishes, the Agnatha, although emerging classification schemes separate them into chordate jawless fishes (the hagfishes) and vertebrate jawless fishes (the lampreys).

Animals that possess jaws are known as gnathostomes, which means “jawed mouth.” Gnathostomes include fishes and tetrapods. Tetrapod literally means “four-footed,” which refers to the phylogenetic history of various land vertebrates, even though in some of the tetrapods, the limbs may have been modified for purposes other than walking. Tetrapods include amphibians, reptiles, birds, and mammals, and technically could also refer to the extinct fishlike groups that gave rise to the tetrapods. Tetrapods can be further divided into two groups: amphibians and amniotes. Amniotes are animals whose eggs contain four extraembryonic membranes (yolk sac, amnion, chorion, and allantois) that provide nutrition and a water-retaining environment for their embryos. Amniotes are adapted for terrestrial living, and include mammals, reptiles, and birds.

Figure 29.2 Deuterostome phylogeny. All chordates are deuterostomes possessing a notochord at some stage of their life cycle.

 

Footnotes

• 1Chen, J. Y., Huang, D. Y., and Li, C. W., “An early Cambrian craniate-like chordate,” Nature 402 (1999): 518–522, doi:10.1038/990080.

 

---

This page titled 13.3: Evolution of Vertebrata/Craniata is a derivative of Biology 2e by OpenStax that is licensed under a CC BY 4.0 license.