Effects of an Epidermal Growth Factor on Growth of Zea Primary Roots and Mesocotyls

Sections of the elongation zones of primary roots and mesocotylsof Zea mays L. were incubated with various concentrations ofhuman epidermal growth factor (EGF). The growth rates of rootsections incubated with 1,000, 100 and 10 µg liter^–1EGF were higher than that of control by 15%, 26% and 14%, respectively.The rates of mesocotyl sections incubated with 100, 10, 1.0and 0.1µg liter^–1 EGF were 23%, 31%, 24% and 22%higher than that of control. (Received August 8, 1994; Accepted November 1, 1994)     

Epidermal Growth Factor Interacts with Indole-3-Acetic Acid and Promotes Coleoptile Growth

We used coleoptile sections of Avena sativa, Sorghum bicolor,and Zea mays seedlings to examine interactions between epidermalgrowth factor (EGF) and indole-3-acetic acid (IAA) that mayaffect plant growth and development. Our 24-h bioassays employedthree controls ranging in dilution from 10^–4 to 10^–8g ml^–1: (1) 50 mM potassium-phosphate buffer solution(pH=6.0), (2) bovine serum albumin, a nonspecific protein; and(3) IAA; plus two treatments: (1) mouse epidermal growth factor(EGF) ranging from 10^–6 to 10^–10gml^–1, and(2) EGF + IAA. In all three species growth in IAA, EGF, andEGF + IAA treatments showed significant increases over controls;EGF+IAA showed significant increases in growth over IAA alone.As the concentrations of IAA decreased, the EGF and IAA interactionbecame more pronounced. At the highest IAA concentrations, EGF+ IAA increased growth rates ca. 2% to 39%, whereas at lowerIAA concentrations EGF + IAA promoted growth as much as 121%,thereby lowering the normal IAA physiological set point up tothree or four orders of magnitude. Our data suggest that aninteraction between EGF and IAA may allow plants to recognizeand respond to animal biochemical messengers, resulting in changesin plant cell elongation that ultimately may alter plant growthpatterns. (Received April 27, 1994; Accepted September 5, 1994)     

Spatial and Temporal Aspects of Growth in the Primary Root of Cotton Seedlings: Effects of NaCl and CaCl2

Seedlings of cotton (Gossypium hirsutum L. cv. Acala SJ-2) weregrown in modified Hoagland nutrient solution with various combinationsof NaCl and CaCl₂. Marking experiments and numerical analysiswere conducted to characterize the spatial and temporal patternsof cotton root growth at varied Na/Ca ratios. At 1 mol m^–3Ca, 150 mol m^–3 NaCl reduced overall root elongation rateto 60% of the control, while increasing Ca to 10 mol m^–3at the same NaCl concentration restored the elongation rateto 80% of the control. Analysis of the spatial distributionof elongation revealed that the presence of 150 mol m^–3NaCl in the medium shortened the growth zone by about 2 mm fromthe approximate 10 mm in the control and also reduced the relativeelemental elongation rate (i.e. the longitudinal strain rate,defined as the derivatives of displacement velocity of a cellularparticle with respect to position on root axis). Supply of 10mol m^–3 Ca at the high salt condition restored partiallythe relative elemental elongation rate, but not the length ofthe growth zone. Compared to the control, the growth trajectoriesshowed that at 1 mol m^–3 CaCl2 it took more time for acellular particle to move through the growth zone at 150 molm^–3 NaCl, while at 10 mol m^–3 CaCl it took lesstime and there was no difference between the NaCl treatments Key words: Gossypium hirsutum, salinity stress, root growth kinematics     

Growth factor-mediated K+ channel activity associated with human myeloblastic ML-1 cell proliferation

ML-1 cell proliferation is dependent on the presence of serumgrowth factors. Removing serum from the culture medium results ingrowth arrest and promotes differentiation. In this study, we foundthat a 4-aminopyridine-sensitiveK⁺ channel was highly expressed inproliferating ML-1 cells and significantly diminished inG₁-arrested ML-1 cells induced by serum deprivation but was restored within 30 min in these cells withaddition of 10% fetal bovine serum (FBS) or 5 ng/ml epidermal growthfactor (EGF). Intracellular adenosine 3',5'-cyclicmonophosphate (cAMP) levels, but not guanosine 3',5'-cyclicmonophosphate, were significantly increased in serum-deprived cellsstimulated by FBS or EGF, and the effects of FBS and EGF on the channelactivation were mimicked by exogenous cAMP. In inside-out patches,K⁺ channel activity wassignificantly increased by the cAMP-dependent protein kinase catalyticsubunit, whereas the effect of EGF on K⁺ channel activation was blockedby Rp-8-(4-chlorophenylthio)adenosine 3',5'-cyclic monophosphothioate. Together, our resultsdemonstrate that serum growth factors stimulateK⁺ channel activity inproliferation of ML-1 cells through protein kinase-inducedphosphorylation and suggest an important molecular mechanism for serumgrowth factor-stimulated mitogenesis in ML-1 cells.

     

Promotion of Plant Cell Division by an Epidermal Growth Factor

In some specified treatments, an epidermal growth factor (EGF)promoted adventitious root formation in epicotyl cuttings ofVigna angularis. The number of the roots induced in cuttingstreated with 0.1 mg liter^-1 EGF during the first 24 h and with210^-4 M IAA during the second 24 h was 15% greater than thatof the roots in cuttings treated without EGF and with IAA. Analysisof the optimum timing of EGF application was performed by dividingthe first 24 h period into three sequential 8 h periods (0–8h, 8–16 h and 16–24 h). The most effective timeperiods in terms of the root formation were 8–16 h and16–24 h. The 0–8 h period was ineffective with respectto the formation. When carrot suspension cells were culturedfor 15 days at a very low cell density (1,000 cells/3 ml Murashigeand Skoog's medium) with more than 0.1 mg liter^-1 EGF, cellnumbers were 72% higher than those cultured without EGF. Theseresults suggest that EGF promotes cell division of plants. (Received October 5, 1992; Accepted May 24, 1993)     

Control of Seed Growth in Soya Beans [Glycine max (L.) Merrill]

The seed is the primary sink for photosynthate during reproductivegrowth and an understanding of the mechanisms controlling therate of seed growth is necessary to understand completely theyield production process. The growth rate of individual seedsof seven soya bean [Glycine max (L.) Merrill] cultivars withgenetic differences in seed size varied from 10.8 to 3.9 mgseed^–1 day^–1. The growth rates were highly correlatedwith final seed size. The growth rate of cotyledons culturedin a complete nutrient medium was highly correlated with thegrowth rate of seeds developing on the plant and with finalseed size. The number of cells per seed in the cotyledons variedfrom 10.2 to 5.7 x 10⁶ across the seven cultivars. The numberof cells per seed in the cotyledons was significantly correlatedwith final seed size and the seed growth rate both on the plantand in the culture medium. The data suggest that genetic differencesin seed growth rates are controlled by the cotyledons and thenumber of cells in the cotyledons may be the mechanism of control. Glycine max L., soya bean, seed size, growth rate, cell number, sink activity     

Growth and production of heterotrophic nanoflagellates in a meso-eutrophic lake

The growth of heterotrophic nanoflagellates (HNF) in mesotrophicLake Constance was measured in situ during a 13 month period.Experiments were conducted with 10 µm pre-filtered lakewater incubated in diffusion chambers at 3 m water depth atthe sampling location for 24 h. Growth rates were calculatedfrom changes in cell numbers occurring during the period ofincubation. Growth rates of all dominant taxa showed pronouncedseasonal variation (–0.13 to 1.76 day^–1 and weregenerally highest in summer at high water temperatures. In situgrowth rates were well below maximum growth rates known forthe respective and similar species from laboratory experiments.While water temperature was a key parameter positively relatedto the growth of all HNF species, the effect of various potentialfood items was taxon specific and less clear. Bacterial abundancewas equally important as temperature for growth in the smallbactenvorous Spumella sp., but was insignificant for growthrates of the larger omnivorous Kathablepharis sp. In Spuniellasp., 84% of the observed seasonal variation of its growth ratecould be explained by temperature and bacterial food supply.Based on these results, a multiple linear regression equationwith temperature and bacterial concentration as dependent variableswas calculated for the growth rate of Spumella. Taxon-specificproduction rates were derived from growth rates and averagebiomass of these two species, and compared to total HNF productionestimated from previously measured community growth rates andbiomass in Lake Constance. Production peaks of Spumella sp.and Kathablepharis sp. alternated seasonally. Total HINF productionranged from –0.01 to 10 mg C m^–3 day^–1. Theaverage seasonal production varied between 1.4 and 33 mg C m^–3day^–1 over 6 consecutive years. These small protozoa thuscontribute a substantial amount to total zooplankton productionin Lake Constance.     

Influence of Cadmium on Growth and Nitrogen Metabolism of Anabaena cylindrica Lemm

A study was made of the effects of cadmium on the Cyanobacterium(blue-green alga) Anabaena cylindrica Lemm. as part of the paddy-fieldecosystem. A simple culture vessel has been designed, which allows periodicalmeasurement of growth (optical density) and nitrogenase activity(C₂H₂-C₂H₄ method). The influence of medium renewal was checked:the renewal of the medium maintained a higher growth rate andhigher nitrogen fixation ability. The cadmium effects were studied using six concentration levelsranging from 0 (control) to 2 parts 10^–6 with renewedmedia (10% every day). No significant differences could be seen up to 1 part 10^–6for nitrogenase activity and relative percentage of heterocysts(decreasing as a function of time from ±4% to ±1.5%). Inhibition of growth (OD and dry weight) was weak at 1 part10^–6 but important at 2 parts 10^–6; at this concentrationcadmium induced morphological and physiological effects: chlorosis,cellular malformations and destruction, and increase in heterocystfrequency (up to 7.72% ±0.19). The cadmium concentration factors were much lower than thosereported for other plants like Chlorella and water pests     

Long-term Respiratory Cost of Maintenance and Growth of Field-grown Young Hinoki Cypress (Chamaecyparis obtusa)

Respiration rate of the entire above-ground parts of field-grown8-year-old hinoki cypress [Chamaecyparis obtusa(Sieb. et Zucc.)Endl.] was measured at monthly intervals over a 5-year period,to evaluate the trend in proportion of maintenance and growthcomponents of respiration with stand development. Representativesample trees were selected for respiration measurements. Theannual respiration rates of individual sample trees were combinedand partitioned into maintenance and growth components by regressingspecific respiration rate on mean relative growth rate. Maintenanceand growth respiration coefficients generated in this way were5.2 mol CO₂kg^-1year^-1and 39 mol  CO₂kg^-1, which are equivalentto 14.3 mg C kg^-1C h^-1(at mean annual air temperature of 15.1°C) and 0.94 kg C kg^-1C. Considering the maintenance andgrowth respiration coefficients, and phytomass and phytomassincrement of individual trees in the stand, the maintenanceand growth respiration rates of the stand were estimated. Theproportion of the maintenance respiration increased, whereasthat of the growth respiration decreased with stand development,due to decreasing relative growth rate.Copyright 1997 Annalsof Botany Company Chamaecyparis obtusa; growth respiration coefficient; hinoki; maintenance respiration coefficient; stand respiration     

Effect of Mechanical Stress on Sunflower (Helianthus annuus L.) Hypocotyl Growth

Etiolated seedlings of Sunflower (Helianthus annuus L.) weresubjected to mechanical stress by longitudinally compressingthe hypocotyl with approx. 0·05 N force (equivalent to0·025 MPa for a 1·6 mm diameter hypocotyl). Thisrelatively low compressive stress effected an increase in relativegrowth rates (RGR) of the hypocotyl for a period of 1-2 h, followingwhich RGR returned to the pretreatment rate. RGR was also increasedby an equivalent compressive stress treatment (0·025MPa) for 4 h in water or in 10^-10 mol l^-1 IAA. These resultsare discussed in the context of a possible role for mechanicallyinduced stress in the initiation and maintenance of nutationalgrowth movements.Copyright 1994, 1999 Academic Press Helianthus annuus, Sunflower, hypocotyl growth, mechanical stress, seedling growth, nutational growth movements, circumnutation     

Effect of Growth Regulators and Light on Pollen Germination and Pollen Tube Growth in Pinus roxburghii Sarg

Pine (Pinus roxburghii) pollen grown in suspension cultureswas used to study the effects of growth regulators and lightconditions on germination and pollen tube growth. Indol-3-ylacetic acid, gibberellic acid, ethylene, abscisic acid and cyclicAMP (cAMP) at low concentrations (1–10 mg 1^–1) promotedgermination and tube growth. Addition of 1 and 10 mg 1^–1cAMP to any of the growth regulators had a promotory effect.Pollen tube growth decreased in white light as compared to thedark, and was increased in red light. Far-red light counteractedthe effect of red light. The effect of growth regulators incausing the enhanced tube growth appears to be manifested throughsubstances such as cAMP, and phytochrome seems to be involved. Pinus roxburghii, pine, pollen germination, pollen tube growth, growth regulators, cyclic AMP, phytochrome     

Effects of Temperature on Growth and Nitrogen Fixation by Swards of Subterranean Clover

Subterranean clover (Trifolium subterraneum L.) cv. ‘Woogenellup’ swards were grown at 10, 15, 20 and 25 Cwith a 12 h photoperiod of 500 or 1000 µmol m^–2s^–1 [low and high photosynthetic photon flux density (PPFD)].Nitrogen-fixing swards received nutrient solution lacking combinednitrogen while control swards received a complete nutrient solution.Growth was measured by infra-red analysis of carbon dioxideexchange and by accumulation of dry matter. Swards were harvestedat intervals between 95 and 570 g d. wt m^–2 for estimationof nitrogenase activity by acetylene reduction and hydrogenevolution assays. Nitrogen fixation was also measured by increasein organic nitrogen. The growth rate was highest at 10 C at low PPFD, and at 10–15C at high PPFD. Nitrogen-fixing swards grew slower than thosereceiving combined nitrogen. Nitrogen fixation measured by increasein organic nitrogen responded similarly to the growth rate,as did acetylene reduction between 10 and 20 C. At 25 C therelationship between acetylene reduction and nitrogen fixationwas distrupted. The difference between the rates of acetylenereduction and hydrogen evolution, theoretically proportionalto nitrogen fixation, was not a reliable indicator of nitrogenfixation because hydrogen uptake developed. Trifolium subterraneum L, subterranean clover, growth, nitrogen fixation, temperature, acetylene reduction     

Modulation of chloride secretory responses and barrier function of intestinal epithelial cells by the Salmonella effector protein SigD

The Salmonella effector protein SigD is an inositol phosphate phosphatase that inhibits phosphatidylinositol 3-kinase-dependent signaling. Because epidermal growth factor (EGF) inhibits chloride secretion via phosphatidylinositol 3-kinase, we explored whether Salmonella infection might modify the inhibitory effect of EGF. As expected, EGF inhibited chloride secretion induced by carbachol in T₈₄ epithelial cells. Infection with wild-type (WT) but not sigD^– mutant S. typhimurium SL1344 decreased CCh-stimulated chloride secretion. Moreover, WT but not sigD^– Salmonella reduced the inhibitory effect of EGF on carbachol-stimulated chloride secretion. Complementation of sigD restored the ability of mutant Salmonella to reverse the inhibitory effect of EGF. EGF-induced EGF receptor phosphorylation was similar in cells infected with either WT or mutant Salmonella, and neither WT nor sigD^– Salmonella altered recruitment of the p85 subunit of phosphatidylinositol 3-kinase to EGF receptor, implying that SigD acts downstream of these signaling events. Furthermore, transepithelial resistance fell more rapidly in cells infected with WT vs. sigD^– Salmonella, indicating an early role for SigD in reducing barrier function, perhaps via activation of protein kinase C. We conclude that the Salmonella bacterial effector protein SigD may play critical roles in the pathogenesis of disease caused by this microorganism. chloride secretion; Salmonella typhimurium; epidermal growth factor     

Factors Affecting Growth and Aggregate Dissociation in Batch Suspension Cultures of Datura innoxia (Miller)

Growth kinetics of Datura innoxia batch suspension cultureswen monitored by a Klett-turbidimetric technique. While cultured. wt varied linearly with Klett units, f. wt and packed cellvolume did not. Turbidimetrically determined doubling timeswere highly reproducible. The method proved to be useful inthe determination of acutely lethal conantrations of a seriesof anti-metabolites. In certain circumstances, aggregate dissociation in batch suspensioncultures of D. innoxia was found to be coupled to growth rate.Suspensions maintained with 10^–5 M 2,4-D exhibited a relativelyslow growth rate with a high degree of aggregate dissociation:10^–4 M 2,4-D promoted a maximum growth rate, but dramaticallysuppressed aggregate dissociation. At 10^–5 M 2,4-D, themitotic index of smaller-aggregate fractions was greater thanthe mitotic index of the large-aggregate fraction. At 10^–5M 2,4-D the converse was observed. Supraoptimal 2,4-D concentrationsthus enhanced both aggregate dissociation and the growth ofsmaller aggregates. When present in concentrations promoting optimal growth. malicand succinic acids caused a decrease in aggregate dissociation.Casein hydrolysate dramatically enhanced growth, but did notaffect aggregate dissociation to the same degree as 2,4-D orthe Krebs cycle organic acids. Suggestions are made concerningmedium composition to be used in future mutant selection schemesusing D. innoxia. Datura innoxia (Miller), suspension culture, growth kinetics, mitotic index, 2,4-dichorophenoxy acetic acid     

Differences in the Interacting Effects of Light and Temperature on Growth of Four Species in the Vegetative Phase

A comparative study, employing the concepts of growth analysis,has been made of the varying responses in the early vegetativephase of Gossypium hirsutum, Helianthus annuus, Phaseolus vulgaris,and Zea mays to combinations of light intensity (1.08, 2.16,3.24, 4.32, and 5.4 x 10⁴ lx—photoperiod 14 h) and constantdiurnal air temperatures (10, 15, 20, 25, 30, and 35 °C).Depending on the combination of treatments, the temperatureof the internal tissues departed from air temperature by 6.9to 1.4 °C: so only the internal temperatures are cited here. For each species there are complex interactions between theeffects of light and temperature on the net assimilation rate,the leaf-area ratio, and the relative growth-rates of plantweight and leaf area. The magnitude of the changes induced bythe two factors vary both with the growth component and thespecies. The temperature responses are maximal up to 20–5°C while at the highest temperatures they may be negative.The temperature coefficients for leaf-area ratio are consistentlyless than those of the other three components: here betweenspecies the coefficients over 10–20 °C vary by a factorof 9.6, 5.4, and 5.1 for the rates of gain in plant weight andleaf area and the net assimilation rate, while the orderingwithin each growth component is species dependent. Under conditions of optimal temperature the relative growth-rateand net assimilation rate progressively increase, accordingto the species, up to either 4.32 or 5.4x 10⁴ lx. The leaf-arearatio is always largest at the lowest intensity. The level oflight at which the rate of gain in leaf area reaches a maximumranges from 2.16x 10⁴ lx for Phaseolus to between 4.32 and 5.40x10⁴ lx for Gossypium. The highest relative growth-rate and net assimilation rate ofHelianthus exceed those of Zea substantially. Indeed the maximalassimilation rate for Helianthus of 2.10 g dm^–2 week^–1is the highest ever recorded under field or controlled conditions.Possible reasons for this reversal of the photosynthetic potentialsof the two species observed by previous workers are discussed.     

Down-regulation of trypsinogen expression is associated with growth retardation in alpha1,6-fucosyltransferase-deficient mice: attenuation of proteinase-activated receptor 2 activity

Alpha1,6-fucosyltransferase (Fut8) plays important roles inphysiological and pathological conditions. Fut8-deficient (Fut8^–/–)mice exhibit growth retardation, earlier postnatal death, andemphysema-like phenotype. To investigate the underlying molecularmechanism by which growth retardation occurs, we examined themRNA expression levels of Fut8^–/– embryos (18.5days postcoitum [dpc]) using a cDNA microarray. The DNA microarrayand real-time polymerase chain reaction (PCR) analysis showedthat a group of genes, including trypsinogens 4, 7, 8, 11, 16,and 20, were down-regulated in Fut8^–/– embryos.Consistently, the expression of trypsinogen proteins was foundto be lower in Fut8^–/– mice in the duodenum, smallintestine, and pancreas. Trypsin, an active form of trypsinogen,regulates cell growth through a G-protein-coupled receptor,the proteinase-activated receptor 2 (PAR-2). In a cell culturesystem, a Fut8 knockdown mouse pancreatic acinar cell carcinoma,TGP49-Fut8-KDs, showed decreased growth rate, similar to thatseen in Fut8^–/– mice, and the decreased growth ratewas rescued by the application of the PAR-2-activating peptide(SLIGRL-NH₂). Moreover, epidermal growth factor (EGF)-inducedreceptor phosphorylation was attenuated in TGP49-Fut8-KDs, whichwas highly associated with a reduction of trypsinogens mRNAlevels. The addition of exogenous EGF recovered c-fos, c-jun,and trypsinogen mRNA expression in TGP49-Fut8-KDs. Again, theEGF-induced up-regulation of c-fos and c-jun mRNA expressionwas significantly blocked by the protein kinase C (PKC) inhibitor.Our findings clearly demonstrate a relationship between Fut8and the regulation of EGF receptor (EGFR)-trypsin-PAR-2 pathwayin controlling cell growth and that the EGFR-trypsin-PAR-2 pathwayis suppressed in TGP49-Fut8-KDs as well as in Fut8^–/–mice.     

Soya Bean Seed Growth and Maturation In vitro without Pods

Immature Glycine max (L.) Merrill seeds, initially between 50and 450 mg f. wt, were grown and matured successfully in vitro.Excised seeds were floated in a liquid medium containing 5 percent sucrose, minerals and glutamine in flasks incubated at25 °C under 300 to 350 µE m^–2 s^–1 fluorescentlight. During 16 to 21 d in culture, seeds grew to a matured. wt of 100 to 600 mg per seed at an average rate of 5 to 25mg d. wt per seed d^–1 depending on initial size. Growthrates were maximal during the first 8 to 10 d in vitro but declinedwith loss of green colour in the cotyledons. Seed coats rupturedwith rapid cotyledon expansion during the first 2 d in culture.Embryos were tolerant to desiccation and 80 to 90 per cent germinatedif removed from culture before complete loss of green colour.The growth of excised seeds in vitro exceeded the growth ofseeds in detached pods, but when windows were cut in pods topermit direct exposure of seeds to the medium, seed growth wascomparable. Glycine max (L.) Merrill, soya bean, seed culture, seed growth, seed maturation, germination     

Effect of Indoleacetic Acid on Growth and Dinitrogen Fixation in Cyanobacteria

The effect of IAA on growth, dinitrogen fixation, and heterocystsfrequency of Anabaena PCC 7119 and Nodularia sp. have been investigated.Concentrations of IAA ranging from 10^–10 to 10^–4M did not change the growth of Anabaena PCC 7119. Concentrationshigher than 10^–4 M were inhibitory. Similar results werefound in Nodularia sp. although in this case the inhibitoryeffect appeared with 10^–5M of IAA. Neither the nitrogenaseactivity nor the heterocysts frequency were enhanced by IAAtreatment. (Received June 17, 1986; Accepted January 22, 1987)     

Effect of Spermidine and Methylglyoxal-bis (Guanyl-hydrazone) (MGBG) on In Vitro Pollen Germination and Tube Growth in Catharanthus roseus

Incorporation of polyamine-spermidine into the nutrient mediumat 10^–6 and 10^–5 M concentrations stimulates pollen-tubegrowth in vitro in Catharanthus roseus L. G. Don. MGBG, an inhibitorof spermidine biosynthesis, at 0.5 x 10^–3 and 1 x 10^–3M concentrations reduced the percentage of germination as wellas tube growth and at a concentration of 1.5 x 10^–3 Mgermination was totally inhibited. Pollen grains incubated inthe medium containing 1.5 x 10^–3 M MGBG, when transferredto a fresh medium with 10^–5 M spermidine, resulted in80% germination recovery, along with considerable tube growth.Experiments with actinomycin-D indicate that stimulation ofpollen-tube growth by spermidine may involve de novo synthesisof protein. Catharanthus roseus, pollen germination, tube growth, spermidine, MGBG, inhibition, actinomycin-D     

Leaf Growth in Cotton (Gossypium hirsutum L.) 1. Growth in Area of Main-stem and Sympodial Leaves

A model is presented for growth of individual and successivemain-stem leaves of cotton, based on a series of indoor experimentsand data sets from the literature. Three variable parametersare used to describe individual leaf growth: relative growthrate of meristematic tissue (R¹), relative rate of approachof final area (R₂) and a ‘position parameter’ (t_0.5)which governs the transition from meristematic to extensiongrowth. Final area of a leaf does not occur in the model asa deterministic quantity but it is a result of the processesduring growth. The model generates successive mainstem leavesand sympodial leaves as an integrated system. Assimilate shortagesoccurring in the plant operate on R₁ leading to the characteristicchange of final leaf area along the mainstem. Gossypium hirsutumL., cotton, leaf growth, relative growth rate, meristematic tissue, extension growth, mathematical model