RESUMEN
The hypothesis of this experiment was that dietary fructose would influence visceral organ mass, carbohydrase activity, and mRNA expression of carbohydrases and nutrient transporters in the small intestine in neonatal calves. Therefore, our objective was to use the neonatal calf as a model to evaluate the effects of postruminal fructose supply on small intestinal carbohydrate assimilation. Ten calves (<7 d of age; 41.2 ± 1.46 kg of body weight) were fed milk replacer at 2.0% of body weight daily (816 ± 90.5 g/d; 272 ± 30.1 g/L; dry-matter basis) in 2 equal portions and assigned to the following dietary treatment groups: (1) milk replacer (control; n = 6) or (2) milk replacer + 2.2 g of fructose/kg of body weight (fructose; n = 4). Calves were fed dietary treatments for 28 d, with jugular blood sampled every 7 d before and after the morning feeding. Calves were slaughtered, and visceral weights were recorded. Postruminal carbohydrase activities were assayed. Quantitative real-time PCR was conducted for small intestinal mRNA expression of nutrient transporters [solute carrier family 2 member 5 (GLUT5), solute carrier family 2 member 2 (GLUT2), and solute carrier family 5 member 1 (SGLT1)], carbohydrases (lactase, maltase-glucoamylase, and sucrase-isomaltase), and ketohexokinase (KHK). Data were analyzed using MIXED procedures in SAS version 9.4 (SAS Institute Inc, Cary, NC). Dietary fructose supplementation decreased serum glucose concentration. Small intestinal mass was greater in calves supplemented with fructose. Dietary fructose supplementation did not influence pancreatic α-amylase, small intestinal isomaltase, or maltase activities. Sucrase activity was undetected in the small intestine. Dietary fructose supplementation increased small intestinal glucoamylase activity per gram of tissue by 30% and increased maltase-glucoamylase mRNA expression by 6.8-fold. Dietary fructose supplementation did not influence mRNA expression of GLUT5, SGLT1, GLUT2, or KHK. Dietary fructose supplementation increased small intestinal lactase mRNA expression by 3.1-fold. Sucrase-isomaltase mRNA expression in the small intestine decreased 5.1-fold with dietary fructose supplementation. Dietary fructose supplementation does not induce sucrase activity in neonatal calves; however, sucrase-isomaltase may be transcriptionally regulated by dietary fructose in neonatal calves. More research is needed to compare glucose and fructose at isocaloric intakes to examine effects of dietary fructose at equal metabolizable energy intake.
Asunto(s)
Metabolismo de los Hidratos de Carbono/genética , Bovinos/metabolismo , Suplementos Dietéticos/análisis , Fructosa/farmacología , Glicósido Hidrolasas/metabolismo , Animales , Animales Recién Nacidos , Dieta/veterinaria , Glucosa/metabolismo , Glicósido Hidrolasas/genética , Intestino Delgado/metabolismo , Sustitutos de la Leche/metabolismo , Nutrientes/metabolismo , ARN Mensajero/genéticaRESUMEN
The objective of this project was to develop and test a web-based virtual dairy herd to help students understand the structure and functioning of a dairy herd, and to promote active learning. At the beginning of the course, the instructor defines the profiles of herds to be assigned to students (e.g., herd size, production, diets, fertility). Each student has a unique herd and engages in decision-making for desired management practices in the herd. Modeled events are based on cow physiology and normal dairy herd management practices. Students' activities and decisions include heat detection, insemination, pregnancy diagnosis, dry-off, diet specifications, feeding groups, colostrum and milk-replacer feeding, weaning, treatment of diseases, and milk withdrawal from the tank if antibiotics are used, among others. The daily output provides information on technical indexes, economic performance, counters of incorrect decisions as feedback for students, and score. Time in class can be devoted to discussions of dairy management issues. Additional exercises based on students' own herds (e.g., calculating required space for cows, land for forage production, manure management) can also be implemented. Students' performance in the virtual dairy farm was monitored over 3 years. The average score (n = 326) was 87.8 ± 1.1 over 100 points, suggesting that self-learning with the virtual dairy farm was highly successful. At the end of each semester, students (n = 277) responded to a survey on the experience of working with the virtual dairy herd. Most students (>87%) agreed that the virtual dairy herd was more effective and motivating than traditional lectures and helped them understand dairy production better. In an unannounced test conducted at least 2 wk before the final exam, students (n = 207) were asked 14 questions on dairy cattle and 14 similar questions on other species taught in the same class through traditional lectures. A similar test on the same students (n = 142) was conducted in their fifth semester (2 years later). Results were better in dairy compared with other species questions in the first (9.6 vs. 3.7) and fifth (8.0 vs. 3.8) semesters. The virtual dairy herd is an effective tool for teaching introductory courses in dairy production. The program can be accessed at www.virtualdairyfarm.org, and a manual and videos with instructions for instructors and students are available online.
Asunto(s)
Bovinos/fisiología , Calostro/metabolismo , Industria Lechera/educación , Fertilidad , Leche/metabolismo , Interfaz Usuario-Computador , Animales , Dieta/veterinaria , Femenino , Inseminación , Lactancia , Sustitutos de la Leche/metabolismo , EmbarazoRESUMEN
The objective of this study was to evaluate the effect of feeding a prepartum diet with a high or moderate starch content on growth and insulin sensitivity of female offspring early in life. Thirty-eight Holstein heifer calves were born to dams fed either a high-starch (26% starch on a DM basis, HI; n = 20) or moderate-starch (14% starch on a DM basis, MOD; n = 18) prepartum diet commencing at 28 ± 3 d before expected parturition date. Following birth, all calves were housed individually and fed three 2-L meals of colostrum within the first 24 h of life and offered 10 L/d of milk replacer (26% CP, 18% fat, mixed to 130 g/L). Body weight of calves was measured at birth and on d 2 (after colostrum feeding but before milk feeding), 10 ± 2, and 20 ± 2. A glucose tolerance test was performed at a minimum of 6 h after their last colostrum or milk meal to evaluate insulin sensitivity on d 2, 10 ± 2 and 20 ± 2. Body weight did not differ throughout between HI and MOD calves; however, calves born to primiparous dams were smaller compared with those born to multiparous dams. Glucose or insulin concentrations were not different before the glucose tolerance test. Following the glucose tolerance test, maximum glucose concentrations were not different between treatments at any time point. However, HI calves had greater insulin area under the curve, and HI calves had greater maximum insulin concentrations on d 2. Glucose or insulin clearance rates were not different nor was the calculated insulin sensitivity index between treatments. These findings suggest that feeding a HI prepartum diet may reduce some insulin sensitivity indicators of female offspring early in life.
Asunto(s)
Alimentación Animal/análisis , Bovinos/fisiología , Calostro/metabolismo , Almidón/administración & dosificación , Animales , Animales Recién Nacidos , Glucemia/análisis , Peso Corporal , Bovinos/crecimiento & desarrollo , Industria Lechera , Dieta/veterinaria , Femenino , Prueba de Tolerancia a la Glucosa/veterinaria , Insulina/sangre , Resistencia a la Insulina , Leche/metabolismo , Sustitutos de la Leche/metabolismo , Parto , EmbarazoRESUMEN
This experiment aimed to evaluate the effects of supplying 4 different inclusion levels of Met + Cys to crossbred liquid-fed calves on animal performance and body composition. Thirty-six Holstein-Gyr male calves were separated into 2 age groups: 16 calves, slaughtered at an age of 30 d, representing the physiological phase from 8 to 30 d, and 20 calves, slaughtered at an age of 60 d, representing the physiological phase from 30 to 60 d. At 8 d of age, the animals were randomly distributed among the experimental treatments: 4 Met + Cys inclusion levels (Met + Cys: 8.0, 8.7, 9.4, and 10.2 g/d), provided by an AA supplement added to 1.0 kg (as fed) of commercial milk replacer containing soy protein concentrate and wheat protein isolate reconstituted at 13.8% (dry matter basis). The diet was supplied without allowing leftovers and no starter feed was provided. The experimental diets were supplied without allowing orts, so that the dry matter, crude protein, and ether extract intakes were the same for all animals, independent of Met + Cys level. Total weight gain, average daily gain, gain composition, and body composition were evaluated for both age groups separately. Digestibility of organic matter, crude protein, and ether extract was lower for 8 to 30 d than for 30 to 60 d. The effect of Met + Cys levels on the digestibility of nutrients was not observed; there also was no significant interaction between physiological phase and Met + Cys levels. For the 8 to 30 d group, no responses in performance were observed according to the different Met + Cys levels, which indicates that 8.0 g/d of Met + Cys met the requirements for this physiological phase. The 30 to 60 d group responded positively to higher Met + Cys inclusion in the diet. In conclusion, an optimal Met + Cys dietary level to ensure best performance and protein gain ranges from 8.41 to 9.81 g/d.
Asunto(s)
Composición Corporal/efectos de los fármacos , Bovinos/crecimiento & desarrollo , Cisteína/farmacología , Metionina/farmacología , Alimentación Animal , Animales , Composición Corporal/fisiología , Cisteína/administración & dosificación , Dieta , Masculino , Metionina/administración & dosificación , Leche , Sustitutos de la Leche/metabolismo , DesteteRESUMEN
Ad libitum milk feeding and butyrate (B) supplementation have the potential to stimulate postnatal growth and development in calves. The somatotropic axis is the main endocrine regulator of postnatal growth and may be affected by both ad libitum milk replacer (MR) feeding and B supplementation in calves. We hypothesized that ad libitum MR feeding and B supplementation stimulate systemic and hepatic insulin-like growth factor (IGF)-I and IGF binding proteins (IGFBP) in preweaning calves. Sixty-four (32 male, 32 female) Holstein calves were examined from birth until wk 11 of life. Calves received MR either ad libitum (Adl) or restrictively (6 L/d; Res). In each feeding group half of the calves received a MR with 0.24% butyrate and the other half received same MR without butyrate. Ad libitum MR feeding was performed from d 4 until wk 8 of age. From wk 9 to 10, Adl and Res calves were gradually weaned and were fed 2 L/d until the end of the trial. Concentrate, hay, and water were freely available. Feed intake was measured daily and body weight weekly. Blood samples for analyzing plasma concentrations of glucose, insulin, IGF-I, and IGFBP-2, -3, and -4 were taken on d 1, 2, 4, and 7, then weekly or every other week (IGFBP) until wk 11 of life. Liver samples were taken on d 50 and at the end of the study (d 80) to measure gene expression of the growth hormone receptor 1A (GHR1A), IGF1, IGFBP1 to 4, and of the IGF Type 1 and insulin receptor in the liver. Intake of MR and body weight were greater, but concentrate intake was lower in Adl than in Res. Plasma concentrations of IGF-I and IGFBP-3 were greater and plasma concentration of IGFBP-2 was lower in Adl than in Res during the ad libitum milk feeding period. After reduction of MR in both groups to 2 L/d plasma concentrations of IGF-I and IGFBP-4 were lower and plasma concentration of IGFBP-2 was higher in Adl than in Res. Supplementation of B depressed plasma IGF-I from wk 1 to 4 and in wk 9. On d 50, mRNA abundance of the GHR1A and IGF1 was greater and of IGFBP2 mRNA was lower in Adl than in Res. At d 80, IGFBP2 mRNA was greater in Adl than in Res, and IGFBP2 mRNA increased with B supplementation. Ad libitum MR feeding stimulated the systemic and hepatic IGF system and mirrored the greater growth rate during the ad libitum MR feeding, whereas butyrate supplementation partly reduced the systemic and hepatic IGF system.
Asunto(s)
Ácido Butírico/administración & dosificación , Bovinos/sangre , Proteínas de Unión a Factor de Crecimiento Similar a la Insulina/metabolismo , Factor I del Crecimiento Similar a la Insulina/metabolismo , Sustitutos de la Leche/administración & dosificación , Alimentación Animal/análisis , Fenómenos Fisiológicos Nutricionales de los Animales , Animales , Ácido Butírico/metabolismo , Bovinos/fisiología , Dieta/veterinaria , Suplementos Dietéticos , Femenino , Regulación de la Expresión Génica/efectos de los fármacos , Insulina/sangre , Proteínas de Unión a Factor de Crecimiento Similar a la Insulina/genética , Factor I del Crecimiento Similar a la Insulina/genética , Hígado/metabolismo , Masculino , Leche/metabolismo , Sustitutos de la Leche/metabolismoRESUMEN
The enhanced growth performance of calves fed a higher plane of nutrition pre-weaning is well documented, and the effect of butyrate on the development of the gastrointestinal tract in calves has been evaluated. The aim of this study was to examine the synergistic effects of ad libitum milk replacer (MR) feeding and butyrate supplementation on growth performance and energy metabolism in calves. Sixty-four (32 male, 32 female) Holstein calves were examined from birth until wk 11 of life. Calves received MR either ad libitum (Adl) or restrictively (Res) with (AdlB+, ResB+) or without (AdlB-, ResB-) 0.24% butyrate supplementation. Colostrum and transition milk were fed in predefined amounts (Res or Adl) for the first 3 d postpartum. Ad libitum and restrictive MR feeding with or without butyrate was performed from d 4 until wk 8 of age. From wk 9 to 10, all calves were gradually weaned and were fed 2 L/d until the end of the trial. Concentrate (CON), hay, and water were freely available. Intakes of MR and CON were measured daily. Calves were weighed at birth and weekly thereafter. Blood was drawn on d 1 before the first colostrum intake; on d 2, 4, and 7; and weekly thereafter until the end of the study to measure plasma concentrations of metabolites and hormones. Liver samples were taken at d 50 and at the end of the study to determine gene expression related to glucose metabolism. Milk, MR, and total nutrient intake were greater, but CON intake was lower in Adl than in Res calves, resulting in a greater body weight, but partially lower gain to feed ratio in Adl than in Res. Plasma concentrations of glucose and insulin were higher during the ad libitum milk-feeding period, whereas plasma ß-hydroxybutyrate was lower in Adl than in Res. Plasma concentrations of nonesterified fatty acids, lactate, total bilirubin, and cortisol were lower, but triglyceride and cholesterol concentrations were higher in Adl than in Res at specific time points. Feed intake, growth performance, and metabolic and endocrine changes were insignificantly affected by butyrate, and hepatic gene expression of enzymes related to endogenous glucose production was barely influenced by ad libitum MR feeding and butyrate supplementation. Intensive MR feeding indicated greater stimulation of growth and anabolic metabolism, but butyrate supplementation did not further improve postnatal growth or anabolic processes either in intensive or restrictive MR-fed calves.
Asunto(s)
Bovinos/crecimiento & desarrollo , Bovinos/metabolismo , Dieta/veterinaria , Sustitutos de la Leche/administración & dosificación , Destete , Ácido 3-Hidroxibutírico/administración & dosificación , Alimentación Animal , Fenómenos Fisiológicos Nutricionales de los Animales , Animales , Animales Recién Nacidos , Calostro , Ingestión de Energía/fisiología , Femenino , Masculino , Leche/metabolismo , Sustitutos de la Leche/metabolismo , EmbarazoRESUMEN
Se reúnen datos de carácter histórico relacionados con los inicios de la introducción y desarrollo de leches maternizadas en nuestro país, que tuvieron lugar durante los años de transición del siglo XIX al XX. En este proceso, destacan de forma muy especial las aportaciones de Luis Soldevila Casas y de los médicos Pau Ferrer Piera i Salvador Soler Farnés. Los productos diseñados y registrados por estos personajes estaban inspirados, básicamente, en los métodos denominados entonces laboratory milk (mezclas) y top milk (fraccionamiento). La vida comercial de estos productos fue muy corta o, incluso, inexistente (AU)
Historic data on the introduction and development of humanized milk that took place in our country during the transition from 19th to 20th century has been compiled. In this process, the contributions of Luis Soldevila Casas and the doctors Pau Ferrer Piera and Salvador Soler Farnés are highly remarkable. The products designed and registered by them were mainly inspired in the processes then known as «Laboratory milk» (blends) and «Top milk» (fractionation). But the shelf life of these products was very short, or they never existed (AU)
Asunto(s)
Humanos , Masculino , Femenino , Recién Nacido , Lactante , Historia del Siglo XIX , Historia del Siglo XX , Lactancia Materna/ética , Lactancia Materna/historia , Lactancia Materna/métodos , Lactancia/fisiología , Sustitutos de la Leche Humana , Sustitutos de la Leche/historia , Sustitutos de la Leche/metabolismo , Leche/fisiología , Lac Vaccinum/uso terapéutico , Pediatría/educación , Pediatría/historia , Pediatría/métodos , Historia de la MedicinaRESUMEN
To clarify colonic fermentation during the perinatal period, 22 dairy calves less than 6 weeks old were used. They were given a milk replacer following colostrum feeding. A total 100 samples of normal feces including meconium were collected from the rectum of the calves. Fecal pH, alcohols, lactate and volatile fatty acids (VFAs) were analyzed. Higher ethanol and n-propanol concentrations were found in many fecal samples particularly in the first 2 weeks after birth, but these metabolites showed consistently lower concentrations thereafter. By contrast, higher concentrations of methanol were observed in some samples for all ages examined. Fecal VFA increased abruptly within a few days of birth, and mainly consisted of acetate and n-butyrate. During the first 2 weeks, the proportion of n-butyrate in VFAs decreased and that of propionate increased gradually. Proportions of VFAs were almost stable at 3-6 weeks of age (acetate, propionate and n-butyrate in increasing order). Higher concentrations of lactate and lower pHs were observed in the fecal samples during the first 2 weeks, and concentrations decreased thereafter. Accelerated colonic production of ethanol and n-propanol was confirmed during the early 2 weeks, in addition to organic acid fermentation as reported previously.
Asunto(s)
1-Propanol/análisis , Animales Recién Nacidos/fisiología , Bovinos , Etanol/análisis , Heces/química , Meconio/química , 1-Propanol/metabolismo , Animales , Aniones/metabolismo , Colon/fisiología , Calostro/metabolismo , Etanol/metabolismo , Ácidos Grasos Volátiles/análisis , Ácidos Grasos Volátiles/metabolismo , Fermentación/fisiología , Concentración de Iones de Hidrógeno , Ácido Láctico/análisis , Ácido Láctico/metabolismo , Sustitutos de la Leche/metabolismoRESUMEN
5'-Uridylic acid (UMP), which is present at high concentrations in cow's colostrum, has been shown to cause a reduction in increased plasma levels of insulin and glucose after ingestion of milk replacer in pre-weaning calves. However, the precise mechanisms of UMP action have not been investigated, and its action has not been investigated in other pre-weaning ruminants. In order to demonstrate whether UMP causes changes in postprandial metabolic and hormonal parameters in pre-weaning goats, 11 Saanen kids were given milk replacer (twice a day) without (n = 5) or with (n = 6) UMP (1 g for each meal, 2 g/day for each head) for 14 days. Analysis of blood samples taken in the morning of day 14 demonstrated that the feeding of milk replacer with UMP abolished the significant changes in postprandial plasma glucose, NEFA, GH and insulin concentrations induced by feeding of milk replacer alone, and demonstrated a tendency to increase IGF-I levels. However, there was no significant difference between the two groups at any sampling time. We conclude that UMP feeding with milk replacer showed a tendency to blunt the postprandial changes in levels of some plasma metabolites and hormones that are induced by replacer alone in pre-weaning goats.
Asunto(s)
Animales Lactantes/metabolismo , Glucemia/metabolismo , Ácidos Grasos no Esterificados/sangre , Hormona del Crecimiento/sangre , Factor I del Crecimiento Similar a la Insulina/metabolismo , Insulina/sangre , Sustitutos de la Leche/metabolismo , Uridina Monofosfato/administración & dosificación , Fenómenos Fisiológicos Nutricionales de los Animales , Animales , Calostro/metabolismo , Cabras , Leche/metabolismo , Periodo Posprandial/fisiología , DesteteRESUMEN
Sixty-four male Holstein-Friesian x Dutch Friesian veal calves (46 +/- 3.0 kg) were used to evaluate the effect of the inclusion of different levels and sources of dietary roughage on animal performance and rumen development. Treatments consisted of 1) C100 = concentrate only; 2) C70-S30 = concentrate (70%) with straw (30%), 3) C70-G30 = concentrate (70%) with dried grass (30%), 4) C70-G15-S15 = concentrate (70%) with dried grass (15%) and straw (15%), 5) C70-CS30 = concentrate (70%) with corn silage (30%), 6) C40-CS60 = concentrate (40%) with corn silage (60%), 7) C70-CS30-AL = concentrate (70%) with corn silage (30%) ad libitum, 8) C70-G15-S15-AL = concentrate (70%) with dried grass (15%) and straw (15%) ad libitum. All dietary treatments were provided in addition to a commercial milk replacer. Concentrate was provided as pellets and roughage was chopped. The dietary treatments 1 to 6 were supplied restrictedly to a maximum of 750 g of dry matter (DM) per day, whereas treatments 7 and 8 were offered ad libitum in combination with a reduced amount of milk replacer. Calves were euthanized after 10 wk. Straw supplementation (C70-S30 vs. C70-G30 and C70-CS30) reduced DM intake, and ad libitum supply of concentrate and roughage increased DM intake. Roughage addition did not affect growth performance. Rumen fermentation was characterized by low pH and high total volatile fatty acids and reducing sugar concentrations. Calves fed ad libitum showed lower ruminal lactate concentrations than calves fed restrictedly. Ammonia concentrations were highest in calves fed C-100 and lowest in calves fed ad libitum. The recovery of CoEDTA (added to milk replacer) varied between 20.5 and 34.9%, indicating that significant amounts of milk entered the rumen. Roughage addition decreased the incidence of plaque formation (rumen mucosa containing focal or multifocal patches with coalescing and adhering papillae covered by a sticky mass of feed, hair and cell debris) and the incidence of calves with poorly developed rumen mucosa. However, morphometric parameters of the rumen wall were hardly influenced by the type and level of roughage. Ruminal polysaccharide-degrading enzyme activities reflected the adaptation of the microorganisms to the dietary concentrate and roughage source. Results indicated that in veal calves, the addition of roughage to concentrate diets did not affect growth performance and positively influenced the macroscopic appearance of the rumen wall.
Asunto(s)
Bovinos/crecimiento & desarrollo , Bovinos/metabolismo , Dieta/veterinaria , Fibras de la Dieta/metabolismo , Sustitutos de la Leche/metabolismo , Rumen/crecimiento & desarrollo , Rumen/metabolismo , Alimentación Animal/análisis , Animales , Ingestión de Alimentos , Ácidos Grasos Volátiles/análisis , Fermentación/fisiología , Contenido Digestivo/química , Contenido Digestivo/enzimología , Concentración de Iones de Hidrógeno , Ácido Láctico/análisis , Masculino , Rumen/anatomía & histología , Factores de Tiempo , Aumento de PesoRESUMEN
This study was designed to investigate the effect of soy protein inclusion in milk replacer diets for goat kids on protein, RNA, and DNA contents in small intestinal mucosa, on the importance of RNA biosynthesis from dietary RNA precursors for mucosal RNA synthesis, and on the activities of enzymes involved in nucleotide degradation in small intestinal mucosa. Diets were based on cow's milk. In the control group, 35% of the milk protein was replaced by casein (CN) protein, and in the soy group (SPAA), the same amount of milk protein was replaced by soy protein supplemented with essential AA known to be at lower concentrations in soy than in CN (Thr, Val, Ile, Leu, His, Lys, Met). Diets were isonitrogenous and isoenergetic. At 47 d of age, goats were harvested and samples of proximal, middle, and distal jejunal mucosa were collected 5 h after feeding 15N-labeled RNA from yeast (13 mg/kg of body weight). Growth and feed conversion did not differ between the control and SPAA kids. Mucosal protein concentrations were lower in the SPAA than the control kids. Concentrations of RNA and DNA did not differ between feeding groups, but in all kids mucosal RNA concentrations were higher in proximal than in middle and distal jejunum. Protein:RNA ratios were higher in the control than the SPAA kids and were lowest in proximal jejunum. Activities of alkaline phosphatase in enterocytes were higher in proximal than in middle and distal jejunum. Activities of mucosal xanthine oxidase were highest in distal jejunum and were higher in the SPAA than the control kids, especially in the middle and distal sites. The 15N-enrichment of mucosal RNA was higher in the control than the SPAA kids, especially in distal jejunum, and was lowest in distal jejunum. In contrast, 15N-enrichment of urea in plasma tended to be higher and Gly concentration in plasma was lower in the SPAA than the control kids. Data indicate that protein content and the protein:RNA ratio were lower in jejunal mucosa of goat kids fed milk replacer with partial replacement of CN protein by soy protein. These findings were accompanied by a lower level of reutilization of preformed dietary RNA precursors for RNA biosynthesis in jejunal mucosa and a higher activity of xanthine oxidase. Thus, feeding soy protein instead of CN protein reduced the incorporation of preformed dietary RNA precursors for RNA biosynthesis in the mucosa and activated key enzymes involved in nucleic acid breakdown.