Figure 20.1. Lungs, which appear as nearly transparent tissue surrounding the heart in this X-ray of a dog (left), are the central organs of the respiratory system. The left lung is smaller than the right lung to accommodate space for the heart. A dog’s nose (right) has a slit on the side of each nostril. When tracking a scent, the slits open, blocking the front of the nostrils. This allows the dog to exhale though the now-open area on the side of the nostrils without losing the scent that is being followed. (credit a: modification of work by Geoff Stearns; credit b: modification of work by Cory Zanker)

Introduction

Breathing is an involuntary event. How often a breath is taken and how much air is inhaled or exhaled are tightly regulated by the respiratory center in the brain. Humans, when they aren’t exerting themselves, breathe approximately 15 times per minute on average. Canines, like the dog in Figure 20.1, have a respiratory rate of about 15–30 breaths per minute. With every inhalation, air fills the lungs, and with every exhalation, air rushes back out. That air is doing more than just inflating and deflating the lungs in the chest cavity. The air contains oxygen that crosses the lung tissue, enters the bloodstream, and travels to organs and tissues. Oxygen (O₂) enters the cells where it is used for metabolic reactions that produce ATP, a high-energy compound. At the same time, these reactions release carbon dioxide (CO₂) as a by-product. CO₂ is toxic and must be eliminated. Carbon dioxide exits the cells, enters the bloodstream, travels back to the lungs, and is expired out of the body during exhalation.

Systems of Gas Exchange

The primary function of the respiratory system is to deliver oxygen to the cells of the body’s tissues and remove carbon dioxide, a cell waste product. The main structures of the human respiratory system are the nasal cavity, the trachea, and lungs.

All aerobic organisms require oxygen to carry out their metabolic functions. Along the evolutionary tree, different organisms have devised different means of obtaining oxygen from the surrounding atmosphere. The environment in which the animal lives greatly determines how an animal respires. The complexity of the respiratory system is correlated with the size of the organism. As animal size increases, diffusion distances increase and the ratio of surface area to volume drops. In unicellular organisms, diffusion across the cell membrane is sufficient for supplying oxygen to the cell (Figure 20.2). Diffusion is a slow, passive transport process. In order for diffusion to be a feasible means of providing oxygen to the cell, the rate of oxygen uptake must match the rate of diffusion across the membrane. In other words, if the cell were very large or thick, diffusion would not be able to provide oxygen quickly enough to the inside of the cell. Therefore, dependence on diffusion as a means of obtaining oxygen and removing carbon dioxide remains feasible only for small organisms or those with highly-flattened bodies, such as many flatworms (Platyhelminthes). Larger organisms had to evolve specialized respiratory tissues, such as gills, lungs, and respiratory passages accompanied by a complex circulatory systems, to transport oxygen throughout their entire body.

Figure 20.2.  The cell of the unicellular algae Ventricaria ventricosa is one of the largest known, reaching one to five centimeters in diameter. Like all single-celled organisms, V. ventricosa exchanges gases across the cell membrane.

Direct Diffusion

For small multicellular organisms, diffusion across the outer membrane is sufficient to meet their oxygen needs. Gas exchange by direct diffusion across surface membranes is efficient for organisms less than 1 mm in diameter. In simple organisms, such as cnidarians and flatworms, every cell in the body is close to the external environment. Their cells are kept moist and gases diffuse quickly via direct diffusion. Flatworms are small, literally flat worms, which ‘breathe’ through diffusion across the outer membrane (Figure 20.3). The flat shape of these organisms increases the surface area for diffusion, ensuring that each cell within the body is close to the outer membrane surface and has access to oxygen. If the flatworm had a cylindrical body, then the cells in the center would not be able to get oxygen.

Figure 20.3.  This flatworm’s process of respiration works by diffusion across the outer membrane. (credit: Stephen Childs)

Skin and Gills

Earthworms and amphibians use their skin (integument) as a respiratory organ. A dense network of capillaries lies just below the skin and facilitates gas exchange between the external environment and the circulatory system. The respiratory surface must be kept moist in order for the gases to dissolve and diffuse across cell membranes.

Organisms that live in water need to obtain oxygen from the water. Oxygen dissolves in water but at a lower concentration than in the atmosphere. The atmosphere has roughly 21 percent oxygen. In water, the oxygen concentration is much smaller than that. Fish and many other aquatic organisms have evolved gills to take up the dissolved oxygen from water (Figure 20.4). Gills are thin tissue filaments that are highly branched and folded. When water passes over the gills, the dissolved oxygen in water rapidly diffuses across the gills into the bloodstream. The circulatory system can then carry the oxygenated blood to the other parts of the body. In animals that contain coelomic fluid instead of blood, oxygen diffuses across the gill surfaces into the coelomic fluid. Gills are found in mollusks, annelids, and crustaceans.

Figure 20.4. This common carp, like many other aquatic organisms, has gills that allow it to obtain oxygen from water. (credit: “Guitardude012″/Wikimedia Commons)

The folded surfaces of the gills provide a large surface area to ensure that the fish gets sufficient oxygen. Diffusion is a process in which material travels from regions of high concentration to low concentration until equilibrium is reached. In this case, blood with a low concentration of oxygen molecules circulates through the gills. The concentration of oxygen molecules in water is higher than the concentration of oxygen molecules in gills. As a result, oxygen molecules diffuse from water (high concentration) to blood (low concentration), as shown in Figure 20.5. Similarly, carbon dioxide molecules in the blood diffuse from the blood (high concentration) to water (low concentration).

Figure 20.5.  As water flows over the gills, oxygen is transferred to blood via the veins. (credit “fish”: modification of work by Duane Raver, NOAA)

Tracheal Systems

Insect respiration is independent of its circulatory system; therefore, the blood does not play a direct role in oxygen transport. Insects have a highly specialized type of respiratory system called the tracheal system, which consists of a network of small tubes that carries oxygen to the entire body. The tracheal system is the most direct and efficient respiratory system in active animals. The tubes in the tracheal system are made of a polymeric material called chitin.

Insect bodies have openings, called spiracles, along the thorax and abdomen. These openings connect to the tubular network, allowing oxygen to pass into the body (Figure 20.6) and regulating the diffusion of CO₂ and water vapor. Air enters and leaves the tracheal system through the spiracles. Some insects can ventilate the tracheal system with body movements.

Figure 20.6.  Insects perform respiration via a tracheal system.

Mammalian Systems

In mammals, pulmonary ventilation occurs via inhalation (breathing). During inhalation, air enters the body through the nasal cavity located just inside the nose (Figure 20.7). As air passes through the nasal cavity, the air is warmed to body temperature and humidified. The respiratory tract is coated with mucus to seal the tissues from direct contact with air. Mucus is high in water. As air crosses these surfaces of the mucous membranes, it picks up water. These processes help equilibrate the air to the body conditions, reducing any damage that cold, dry air can cause. Particulate matter that is floating in the air is removed in the nasal passages via mucus and cilia. The processes of warming, humidifying, and removing particles are important protective mechanisms that prevent damage to the trachea and lungs. Thus, inhalation serves several purposes in addition to bringing oxygen into the respiratory system.

Figure 20.7.  Air enters the respiratory system through the nasal cavity and pharynx, and then passes through the trachea and into the bronchi, which bring air into the lungs. (credit: modification of work by NCI)

Which of the following statements about the mammalian respiratory system is false?

1.      When we breathe in, air travels from the pharynx to the trachea.

2.      The bronchioles branch into bronchi.

3.      Alveolar ducts connect to alveolar sacs.

4.      Gas exchange between the lung and blood takes place in the alveolus.

From the nasal cavity, air passes through the pharynx (throat) and the larynx (voice box), as it makes its way to the trachea (Figure 20.7). The main function of the trachea is to funnel the inhaled air to the lungs and the exhaled air back out of the body. The human trachea is a cylinder about 10 to 12 cm long and 2 cm in diameter that sits in front of the esophagus and extends from the larynx into the chest cavity where it divides into the two primary bronchi at the midthorax. It is made of incomplete rings of hyaline cartilage and smooth muscle (Figure 20.8). The trachea is lined with mucus-producing goblet cells and ciliated epithelia. The cilia propel foreign particles trapped in the mucus toward the pharynx. The cartilage provides strength and support to the trachea to keep the passage open. The smooth muscle can contract, decreasing the trachea’s diameter, which causes expired air to rush upwards from the lungs at a great force. The forced exhalation helps expel mucus when we cough. Smooth muscle can contract or relax, depending on stimuli from the external environment or the body’s nervous system.

Figure 20.8.  The trachea and bronchi are made of incomplete rings of cartilage. (credit: modification of work by Gray’s Anatomy)