Growth Hormone and Its Effects on Growth
Growth Hormone (GH) Experiments
Early 1900s experiments demonstrated that injecting "bovine pituitary extract" into rodents resulted in increased growth (postnatal).
It was observed that:
Removing the pituitary from rodents (hypophysectomy, abbreviated as Hypox) reduced their growth.
Injecting pituitary extract into Hypox rodents restored their growth.
Reference: Hossner 2005 "Hormonal Regulation of Farm Animal Growth".
Significance of Findings
The results indicated that a substance produced by the pituitary gland has major effects on growth.
This substance was later identified as Growth Hormone (GH):
GH is mostly species-specific; injection from other species generally lacks bioactivity.
Exceptions exist if there is a similar structure allowing receptor binding, as seen in monkey and human interactions, and rat and cow experiments.
Characteristics of Growth Hormone (GH)
GH is a protein synthesized by the anterior pituitary gland.
Structural Information:
Composed of approximately 191 amino acids; has a molecular weight of 22 kilodaltons (kd).
Release Patterns:
Released at night in 6-8 pulses, particularly during sleep.
Regulation of Production/Release:
Stimulated by the hypothalamic hormone GHRH (Growth Hormone-Releasing Hormone).
Inhibited by somatostatin (SS).
The GH gene possesses a 5' promoter region for transcription factors.
Major Target Organs for GH
The primary target organ for GH action is the liver, which has a higher number of GH receptors (GHR).
Other target cells include bone, muscle, and adipose tissues, which are also equipped with GHRs.
GH has significant positive effects on muscle and bone tissues, while having a negative effect on adipose tissue.
Effects of GH in Pigs
Recombinant Porcine GH (commercially known as porcine somatotropin, rPST) is utilized:
Administered daily or periodically prior to slaughter.
Results in improved pork quality:
Increased muscle mass.
Decreased fat content.
Interaction of GH and Other Factors
GH does not operate in isolation.
Serum from normal mice can induce radioactive sulfate incorporation into cartilage, indicating cell growth, while serum from Hypox mice cannot.
This implies a missing serum factor that GH interacts with for effective growth.
Discovery of IGF Activity
Insulin-like Growth Factor 1 (IGF-1) discovered as an important component in GH action.
Research by W. H. Daughaday at Washington University, St. Louis.
Function demonstrated by radioactive sulfate incorporation as a marker for growth.
Characteristics of IGF-1
GH exerts its effects through IGF-1.
Comparison with Insulin:
IGF-1 is a single chain peptide consisting of 70 amino acids.
It shares about 50% homology with insulin.
IGF Receptors
IGF-1 binds to its own receptor, creating biological effects, and also binds to the insulin receptor albeit with low affinity.
IGF-II also has a receptor (IGFR2R) with high binding affinity.
The effects of IGF and insulin are interconnected and significant.
Regulation of IGF Effects
The action of IGF is regulated by IGF Binding Proteins (IGFBP-1 through IGFBP-3).
GH stimulates the synthesis of both IGF-1 and IGFBP-3 primarily in the liver, bone, and muscle.
Role of IGF-Binding Proteins (IGFBPs)
IGFBPs modulate the action of IGF via its receptor (IGFR).
Infusion of IGF-1 can induce rapid hypoglycemia, but the presence of IGFBP can mitigate this effect.
IGFBPs are primarily produced in the liver and bind 99% of circulating IGF.
The most prevalent IGFBP, IGFBP3, is synthesized by the liver with over 80% bound to IGF-1, serving as a clinical indicator of IGF-1 activity.
Systemic and Local IGF Effects
The systemic effects of IGF differ from paracrine/autocrine actions.
Paracrine/autocrine IGF generally has a more significant growth impact than endocrine IGF.
Circulating IGF from the liver controls pituitary GH release and regulates local IGF production.
Fetal and Neonate IGF Activity
In late gestation, IGF-1 levels in fetal circulation are correlated with fetal size.
The placenta is the primary source of IGF-1 for the fetus during most of gestation, influencing nutrient transfer.
The fetal liver becomes the major source of IGF-1 in late gestation, and postnatally, IGF-1 levels spike due to GH effects on the liver.
IGF-II Dynamics
IGF-II levels are higher in the fetus, but are not directly related to size; they decline post-birth.
The IGF-II and IGF2R genes are imprinted, expressed monoallelically based on parentage.
For example, in mouse fetuses, only the paternal IGF-II gene is expressed, while only the maternal IGF2R is expressed.
Placental and Fetal IGF Role
IGF is crucial for tissue growth and is produced in response to maternal nutritional factors.
Restrictions during gestation limit fetal and subsequently adult size.
Fetal GH is not necessary for growth, even in the presence of maternal influence.
Example: Fetal hypophysectomy has minimal effect on birth weight in pigs, rats, and sheep.
The placenta provides critical nutritional factors driving fetal IGF production.
Maternal GH-IGF Relationships
Maternal growth is dependent on GH, which is essential for IGF-1 production.
However, maternal GH and IGF do not pass through the placenta to the fetus.
Postnatal Growth Requirements
Postnatal growth hinges on GH from the pituitary and IGF from the liver:
The liver produces IGF due to its high receptor density (GHR).
IGF has major impacts on bone, muscle, and adipose tissues, with these tissues also showcasing GHRs.
Factors Influencing GH Release
GH is released in response to various metabolic and hormonal cues:
Positive Influences: Sleep, nutrition, exercise, and sex hormones.
Negative Influences: Stress and glucocorticoids.
Connection to the gut: When the stomach is empty, it produces Ghrelin, a peptide that stimulates GH release.
Physiological Conditions for Growth
Positive conditions for growth:
Blood glucose levels positive, Insulin positive, Amino acids positive, and GH positive.
Negative conditions for growth include elevated IGF-1 levels, free fatty acids, and suppressed GH.
Mechanisms of Growth Regulation
In the liver, GH enhances IGF-1 production, reduces amino acid oxidation, increases glucose release, and diminishes insulin responsiveness and gluconeogenesis.
In muscle, GH promotes amino acid uptake (enhancing protein synthesis), mobilizes muscle glycogen reserves, and boosts glucose uptake.
In adipose tissues, GH reduces lipogenesis (lowering insulin sensitivity) and promotes lipolysis.
Synergy of IGF and Insulin
Together, IGF and insulin mediate glucose and amino acid uptake into muscle, adipose, and bone tissues via their receptors.
Insulin-dependent transporters for amino acids and glucose become more available on cell surfaces as insulin levels rise.
Insulin's Role in Tissue Metabolism
Insulin is crucial for specific tissues:
Muscle: Utilizes fatty acids and glucose as main fuels; insulin promotes glucose uptake and glycogen storage.
Adipose: Insulin also promotes glucose uptake and fatty acid release.
Brain: Glucose acts as the main fuel (lacks glycogen storage).
Liver: Responsible for glycogen storage and glucose release; uptake is dictated by glucose concentration.
GH and IGF's Effects on Bone Growth
GH before puberty encourages normal long bone growth.
Increased levels of sex steroids (such as estrogen) at puberty halt long bone growth, concluding the growth phase.
Postnatal IGF Observations in Dogs
Studies have shown dogs' postnatal growth in body weight and size parallels IGF-1 levels, with significant findings from Eigenmann, 1984, Acta Endocrinol.
IGF-1 and Height Association in Humans
In humans (both males and females), IGF-1 levels positively correlate with height up until puberty.
This pattern is also observed in rodents, livestock, and domestic cats.
Additionally, breed size in dogs associates with specific IGF-1 alleles.
Implications of Excess GH
In humans and animals before puberty, excess GH can lead to gigantism:
Example: Robert Wadlow (1918-1940), who measured 8'11" due to excessive elongation of bones and continued organ growth, often caused by pituitary tumors.
Consequences of Excess GH After Puberty
Post-puberty excess GH can cause acromegaly:
Example: Andre the Giant, measuring 7'4" and weighing 520 pounds, exhibited enlarged extremities and facial features due to continued bone growth without height increase.
This condition typically relates to pituitary tumors causing elevated GH and IGF-1.
Consequences of Low GH Before Puberty
Insufficient GH results in dwarfism characterized by abnormal proportions due to restricted long bone growth.
The inability to produce or bind GH or IGF can result in a miniature body size, while maintaining normal proportions.
An example includes miniature Brahman cattle that are 70% of expected height, often attributed to pituitary tumors, cellular trauma (such as radiation), or genetic mutations affecting growth pathways.