was important the be grown Nutrient Disorders of ‘Evolution’ under nutrient-stress conditions to Mealy-cup Sage add this information to the grower’s toolbox of diagnostic criterion. To quickly recognize and diagnose nu- Jared Barnes1,3, Brian Whipker1, Ingram McCall1, trient disorders, growers should have and Jonathan Frantz2 knowledge of symptoms and diagnos- tic tools. These diagnostic tools in- clude symptomology descriptions, ADDITIONAL INDEX WORDS. deficiency, macronutrients, , images of the disorders, and refer- farinacea, toxicity ence tissue concentrations. Mealy-cup sage ( fam- SUMMARY. To produce floriculture crops like mealy-cup sage (Salvia farinacea), growers must be equipped with cultural information including the ability to ily) is a bedding annual planted by recognize and characterize nutrient disorders. ‘Evolution’ mealy-cup sage plants gardeners to provide color during the were grown in silica-sand culture to induce, describe, and photograph symptoms of summer (Armitage, 2001). Few nutri- nutritional disorders. Plants received a complete modified Hoagland’s all-nitrate ent disorders have been reported for solution of (macronutrient concentrations in millimoles) 15 nitrate- (N), mealy-cup sage. Interveinal 1.0 (P), 6.0 potassium (K), 5.0 (Ca), 2.0 magnesium (Mg), and was reported as a symptom for Mg 2.0 sulfur (S) plus ( concentrations in micromoles) 72 iron (Fe), 18 deficiency (SePRO, 2000). Symptoms manganese (Mn), 3 copper (Cu), 3 zinc (Zn), 45 boron (B), and 0.1 molybdenum described for high electrical conduc- (Mo). Nutrient-deficient treatments were induced with a complete nutrient formula tivity (EC) included necrotic foliage minus one of the nutrients. The B-toxicity treatment was induced by increasing the (Armitage et al., 1994) and cup- element 10-fold higher than the complete nutrient formula. Reagent-grade chemicals and deionized (DI) of 18 million ohms per centimeter purity were ping (SePRO, 2000). Thick veins on used to formulate treatment solutions. We monitored plants daily to document and young with a gray or brown photograph sequential series of symptoms as they developed. Typical symptomol- coloration was due to high salts or ogy of nutrient disorders and corresponding tissue concentrations were deter- ammonium toxicity (Dreistadt, 2001; mined. Out of 13 treatments, 12 exhibited symptomology; Mo was asymptomatic. SePRO, 2000). Cool temperatures Symptoms of N, P, S, Ca, and K deficiencies and B toxicity manifested early; with reduced nutrition caused chlo- therefore, these disorders may be more likely problems encountered by growers. rosis (SePRO, 2000). Unique symptoms were observed on plants grown under N-, Cu-, and Zn-deficient With the lack of knowledge of conditions. Necrosis was a common symptom observed, but use of other diagnostic other nutritional problems that may criteria about location on the and progression of the disorder can aid growers occur with mealy-cup sage, growers in diagnosing nutrient disorders of mealy-cup sage. may face plant loss, increased cost, and wasted time if a nutritional prob- ach essential element taken up walls, is much more immobile; thus, lem arises in the greenhouse. There- by a plant serves to fulfill a spe- symptoms will typically appear in the fore, the objectives for this research Ecific physiological role, and upper foliage (Gibson et al., 2007). were to determine what nutrient- reduced (nutrient deficiency) or ex- While it is true that across the spec- disorder symptoms occur in mealy- cess (nutrient toxicity) levels of that trum of plant species many similarities cup sage and to elucidate threshold element often result in unique symp- exist in how each nutrient is used, not nutrient levels. tomology that may be used for di- all species will develop symptomology, agnostic purposes. The differences in and some crops may actually man- Materials and methods the specificity of the roles and accom- ifest unique symptomology (Nelson, The experiment was conducted panying nutritional disorders are illus- 2012). For example, Reuther and in a glass greenhouse in Raleigh, NC, trated with N and Ca deficiencies. An Labanauskas (1973) stated a species- at 35N latitude. ‘Evolution’ mealy- important component of amino acids, specific approach is required to de- cup sage plugs (2.7 · 2.7 · 3.6 cm cell proteins, and nucleic acids (Briskin termine symptoms of Cu deficiency. size; Raker, Litchfield, MI) were sin- and Bloom, 2010), N is able to be To evaluate crops for the unique symp- gly transplanted into 5-inch-diameter translocated to actively growing re- toms of species-specific disorders, it (0.76 L vol.) plastic pots containing gions under N-deficient conditions, and symptomology typically manifests in the lower leaves. On the other hand, Ca, an important component of cell Units To convert U.S. to SI, To convert SI to U.S., We are grateful for the funding support provided by multiply by U.S. unit SI unit multiply by USDA-ARS and the Floriculture and Nursery Re- 29.5735 fl oz mL 0.0338 search Initiative, and we would like to thank Benary 0.3048 ft m 3.2808 for the mealy-cup sage seed, and to Doug Sturtz, Russ Friedrich, and Alycia Pittenger for nutrient analysis. 3.7854 gal L 0.2642 2.54 inch(es) cm 0.3937 1 Department of Horticultural Science, North Caro- 25.4 inch(es) mm 0.0394 lina State University, Box 7609, Raleigh, NC, 27695 28.3495 oz g 0.0353 2U.S. Department of Agriculture, Agricultural Re- 28,350 oz mg 3.5274 · 10–5 search Service, Application Technology Research 1 ppm mgÁkg–1 1 Unit, 2801 W. Bancroft, MS 604, Toledo, OH 43606 1 ppm mgÁL–1 1 3Corresponding author. E-mail: [email protected]. (F – 32) O 1.8 F C(1.8·C) + 32

502 • August 2012 22(4) acid-washed silica sand [Millersville diethylenetriaminepentaacetic acid sampling. The remaining five potted #2 (0.8–1.2 mm diameter); Southern (FeDTPA), manganese chloride tet- plants were grown to document ad- Products and Silica Co., Hoffman, rahydrate (MnCl2Á4H2O), zinc chlo- vanced symptoms. Recently matured, NC] on 11 June 2009. Seedlings ride (ZnCl2), cupric chloride dihydrate fully expanded leaves were sampled to were fertilized with each irrigation (CuCl2ÁH2O), boric acid (H3BO3), evaluate the threshold tissue concen- using 150 mgÁL–1 N from 13N– and sodium molybdate dihydrate tration for each element. Harvested 0.9P–10.8K Cal-Mag (SQM North (Na2MoO4Á2H2O). Sodium hydrox- leaves were initially rinsed with DI America, Atlanta). Plants were grown ide (NaOH) was added to adjust the water, then washed in a solution of with 70 F day and 65 F night air pH to 6.0 (Barnes, 2010). 0.5 N HCl for 1 min and again rinsed temperature set points and ambient To induce nutrient-deficient with DI water. The remaining shoot light and natural photoperiods. Be- treatments, the plants were irrigated tissue was harvested separately. Roots cause symptoms of N, P, and Fe with complete nutrient solution ex- were discarded. Both sets of tissue manifested too early to obtain ade- cluding one of the nutrients. For were dried at 70 C for at least 1 week, quate tissue, we grew additional plants macronutrients, because many of the and the dry weights were recorded in complete control and N-, P-, cations and anions listed above were (Table 2). After drying, fully expanded and Fe-deficient conditions to linked to a corresponding required leaf tissue was ground in a sample obtain adequate tissue with the fol- element, we used the salts potassium mill (Foss Tecator Cyclotecä;Ana- lowing exceptions. ‘Evolution’ mealy- chloride (KCl), calcium chloride lytical Instruments, Golden Valley, cup sage seeds were sown into a dihydrate (CaCl2Á2H2O), sodium ni- MN) to pass a £0.5-mm sieve. Tissue 72-plug (4.0 · 4.0 · 5.8 cm cell trate (NaNO3), sodium phosphate analysis for N was performed with dimensions) tray containing Fafard monobasic monohydrate (NaH2- a C–H–N analyzer (model 2400 series 4P (Conrad Fafard, Agawam, MA) PO4ÁH2O), sodium sulfate anhy- II; PerkinElmer, Norwalk, CT) by root substrate. These plants were drous (Na2SO4), and magnesium weighing 3.5 mg of dried tissue into installed in the previously described chloride hexahydrate (MgCl2Á6H2O) tin cups and placing it into the ana- manner on 9 Nov. 2009 into treat- in the stock solutions to supply the lyzer. Other nutrient concentrations ments for control, N-, P-, and Fe- missing element. In each case for were determined with inductively deficient conditions. macronutrients, the salt cations were coupled plasma optical emission spec- Treatments started immediately replaced with sodium (Na), and an- troscopy [(ICP-OES (model IRIS In- upon transplanting. An automated, ions were replaced with chloride (Cl). trepid II; Thermo Corp., Waltham, recirculating irrigation system was For micronutrients, the micronu- MA)]. Tissue concentration values ob- constructed out of 4-inch-diameter trient stock solution containing the tained from this experiment were com- PVC pipe [PVC (Charlotte Plastics, micronutrient salt was omitted from pared with optimum values (Table 3) Charlotte, NC)]. The system con- the solution. The B-toxicity treat- published by Mills and Jones (1996) sisted of 18 separate irrigation lines ment was conducted by increasing B for scarlet sage (Salvia splendens), an- (each 6 ft long). Each line contained concentration to 450 mM in the other bedding sage (Dole and Wilkins, eight openings on the top that con- Hoagland’s solution. Stock solutions 2005). The first experiment for all tained PVC funnels with 5-inch- were added to a 32-gal barrel partially treatments was ended on 28 July diameter openings that held the eight filled with DI water and then topped 2009. At termination any treatments pots for the elemental treatment; 13 up to 100 L using DI water. Irriga- that were symptomless were sampled lines contained the disorder treatments tion was automated using a drip for shoot dry weight and nutrient and five lines contained the control system equipped with sump-pumps concentrations of the recently ex- treatments. Pipes were randomly as- (model 1A; Little Giant Pump Co., panded mature leaves. Additional signed as a control or treatment after Oklahoma City, OK) and plastic plants that were grown for obtain- the system was constructed. Control emitters, and the frequency was ad- ing adequate tissue in the complete plants were grown with a complete justed to maintain plant turgidity so control and under N-, P-, and Fe- modified Hoagland’s all-nitrate solu- that the plants never experienced deficient conditions were ended on tion of (macronutrient concentrations wilting; on average, irrigation oc- 17 Dec. 2009. All the data were sub- inmillimoles)15N,1.0P,6.0K, curred once every 3 h from 0600 to jected to analysis of variance (ANOVA) 5.0 Ca, 2.0 Mg, and 2.0 S (Hoagland 1800 HR. The solution drained out using PROC ANOVA of SAS (version and Arnon, 1950) plus (micronu- from the bottom of the pot into the 9.2; SAS Institute, Cary, NC). Where trient concentrations in micromoles) PVC pipe and was captured for reuse. the F test indicated evidence of sig- 72Fe,18Mn,3Cu,3Zn,45B,and Nutrient solutions were replaced nificant difference among the means, 0.1 Mo. Reagent-grade chemicals weekly. Plants were monitored daily least significant difference at P £ 0.05 and DI water of 18 million ohms to record the date symptoms man- was used to establish differences be- per centimeter purity were used to ifested and to document and photo- tween means. formulate stock solutions (Table 1). graph sequential series of symptoms Nutrients in the stock solutions origi- on youngest, young, recently ma- Results nated from the salts of potassium ture, and/or mature leaves as they Outof13nutrientdisordertreat- nitrate (KNO3), calcium nitrate developed. ments, 12 exhibited symptomology. tetrahydrate [Ca(NO3)2Á4H2O], When the initial symptom oc- Symptoms and shoot dry weight values potassium phosphate monobasic curred on the shoots of each treat- (Table 2) were reported. Values for (KH2PO4), magnesium sulfate ment, the date was recorded and percentage difference in shoot dry heptahydrate (MgSO4Á7H2O), iron three potted plants were selected for weights were presented in the text if

• August 2012 22(4) 503 RESEARCH REPORTS

Table 1. Salts used to formulate stock and base solutions, stock solution molarity, and volume of stock solutions used to prepare 100 L (26.4 gal) of nutrient solution for the mealy-cup sage complete control, nutrient deficiency treatments, and boron toxicity. Stock Volume of stock solution added (mL)z solution Complete Fertilizer salt and base molarity controly LN LP LK LCa LMg LS LB DDB LCu LFe LMn LMo LZn

Potassium nitrate (KNO3) 1 M 500 — 500 — 500 500 500 500 500 500 500 500 500 500 Calcium nitrate tetrahydrate 1 M 500 — 500 500 — 500 500 500 500 500 500 500 500 500 [Ca(NO3)2•4H2O] Potassium phosphate monobasic 1 M 100 100 — — 100 100 100 100 100 100 100 100 100 100 (KH2PO4) Magnesium sulfate heptahydrate 1 M 200 200 200 200 200 — — 200 200 200 200 200 200 200 (MgSO4•7H2O) Potassium chloride (KCl) 1 M — 500 100 — — — — — — — — — — — Calcium chloride dihydrate 1 M — 500——— — —— — —— — — — (CaCl2•2H2O) Sodium nitrate (NaNO3) 1 M — — — 500 500 — — — — — — — — — Sodium phosphate monobasic 1 M — — — 200 — — — — — — — — — — monohydrate (NaH2PO4•H2O) Sodium sulfate anhydrous 1 M — ————200—— — —— — — — (Na2SO4) Magnesium chloride hexahydrate 1 M — — — — — — 200 — — — — — — — (MgCl2•6H2O) Iron diethylenetriaminepentaacetic 1 M 100 100 100 100 100 100 100 100 100 100 — 100 100 100 acid (FeDTPA) Manganese chloride tetrahydrate 20 mM 90 90 90 90 90 90 90 90 90 90 90 — 90 90 (MnCl2•4H2O) Zinc chloride (ZnCl2) 20 mM 15 15 15 15 15 15 15 15 15 15 15 15 15 — Cupric chloride dihydrate 20 mM 15 15 15 15 15 15 15 15 15 — 15 15 15 15 (CuCl2•2H2O) Boric acid (H3BO3) 100 mM 45 45 45 45 45 45 45 — 450 45 45 45 45 45 Sodium molybdate dihydrate 1mM 10 10 10 10 10 10 10 10 10 10 10 10 — 10 (Na2MoO4•2H2O) Sodium hydroxide (NaOH) 1 M 40 40 5 45 40 40 40 6 40 6 6 40 40 40 zTreatment formulations for nitrogen deficiency (LN), phosphorus deficiency (LP), potassium deficiency (LK), calcium deficiency (LCa), magnesium deficiency (LMg), sulfur deficiency (LS), boron deficiency (LB), boron toxicity (++B), copper deficiency (LCu), iron deficiency (LFe), manganese deficiency (LMn), molybdenum deficiency (LMo), and zinc deficiency (LZn); 1 mL = 0.0338 fl oz. yRecipe for the complete modified Hoagland’s all-nitrate solution in which control plants were grown.

Table 2. ‘Evolution’ mealy-cup sage plant dry weight as affected by deficient or toxic nutrient treatments are shown in comparison with the complete control that was harvested at the time symptoms were observed. Dry wt (g)z Element LN LP LK LCa LMg LS LB DDB LCu LFe LMn LMoy LZn Complete control 0.53 1.03 2.33 2.33 2.39 1.04 3.94 2.33 9.91 2.02 6.45 13.99 2.39 Treatment 0.33 0.63 1.45 1.47 2.05 0.58 2.57 2.16 10.45 2.04 5.71 18.44 1.43 P valuex ** * *** *** NS ** NS NS NS NS NS NS * zTreatments for nitrogen deficiency (LN), phosphorus deficiency (LP), potassium deficiency (LK), calcium deficiency (LCa), magnesium deficiency (LMg), sulfur deficiency (LS), boron deficiency (LB), boron toxicity (++B), copper deficiency (LCu), iron deficiency (LFe), manganese deficiency (LMn), molybdenum deficiency (LMo), and zinc deficiency (LZn); 1 g = 0.0353 oz. yAsymptomatic treatment. x*, **, or *** indicate statistically significant differences between sample means based on F test at P £ 0.05, P £ 0.01, or P £ 0.001, respectively; NS indicates the F test difference between sample means was P > 0.05.

control and treatment tissues were NITROGEN. Leaves on plants from plants grown under N-deficient significantly different when visual grown under N-deficient conditions conditions was 2.57%. As the disorder symptoms first appeared (Fig. 1A). became chlorotic and had 38% less progressed, chlorosis intensified, and Unless otherwise noted, values for dry weight than the control plants. the leaf margins developed a purple tissue concentrations presented were The control tissue N concentration coloration (Fig. 1B). Lower leaf tips significantly different (Table 3). was 5.06%, while tissue concentration became necrotic. The necrotic area

504 • August 2012 22(4) Fig. 1. (A) The location where initial symptoms were first observed on ‘Evolution’ mealy-cup sage plants for nitrogen deficiency (LN), phosphorus deficiency (LP), potassium deficiency (LK), calcium deficiency (LCa), magnesium deficiency (LMg), sulfur deficiency (LS), boron deficiency (LB), boron toxicity (DDB),copperdeficiency(LCu), iron deficiency (LFe), manganese deficiency (LMn), molybdenum deficiency (LMo), and zinc deficiency (LZn). (B) Plants grown under nitrogen deficiency exhibited chlorosis. In the advanced stage seen here, a purple coloration was observed on the leaf margins. (C) Plants grown in phosphorus-deficient conditions had less dry weight than controls and exhibited brown necrosis on the lower leaf tips; lower leaves dropped in the advanced stage. (D) Older leaves on potassium-deficient plants developed a ‘‘camouflage’’ necrotic pattern. (E) Upper leaves of plants grown in calcium-deficient conditions exhibited black, necrotic spots on the leaf margin and leaf curl. (F) Middle leaves of plants grown with magnesium deficiency exhibited interveinal gray-brown spots which advanced to interveinal necrosis. (G) A light green coloration manifested in sulfur-deficient plants and intensified in the middle and upper foliage.

Fig. 2. (A) Upper leaves on mealy-cup sage plants growing in boron-deficient conditions developed strap-like leaves and brown spots on the leaf near the petiole. (B) Lower leaves of plants grown under boron toxicity exhibited lower-leaf, marginal necrosis. (C) On plants grown in copper-deficient conditions, green-gray coloration was observed on middle and upper leaves. (D) Plants grown under iron deficiency manifested a light green coloration. (E) Light green coloration progressed to interveinal chlorosis and brown, marginal or interveinal spotting on plants grown under the manganese-deficient regime. (F) Upper leaves on plants grown under zinc deficiency developed a light green coloration that was followed by brown necrotic spots; the inflorescence also exhibited reduced elongation and development.

• August 2012 22(4) 505 RESEARCH REPORTS

advanced inward on the leaf. In the entire leaf became necrotic. The dis-

Zn advanced stage, the upper leaves also coloration then manifested in upper 21.4 12.1 L became necrotic. leaves, and chlorosis began to appear PHOSPHORUS. Deficiency of P in lower leaves. In the advanced stage, = 1 ppm.

–1 manifested on lower leaves with most of the middle leaves then became x w v kg 0.05. Á brown necrotic spots appearing on fully necrotic. Chlorotic, lower leaves > Mo

P the tip of the leaf (Fig. 1C). Plants exhibited some brown discoloration. y L )

1 grown under P-deficient conditions SULFUR. Initially, plants grown L m concentrations Zn); 1 mg had 39% less dry weight than those under conditions of S deficiency de- kg L Á receiving P. Control plants had a con- veloped a light green coloration.

Mn centration of 0.67% P, and plants Plants exhibiting S-deficiency symp- L grown under P-deficient conditions toms had 44% less dry weight than the

S). had a concentration of 0.12% P. As control plants. Tissue concentrations L symptoms progressed, affected leaves were significantly different; control Fe abscised from the plant. Upper leaves tissue was 0.45% S, while the tissue L then developed the necrotic marginal of plants grown without S had 0.24% Mo), and zinc deficiency ( spotting. S. The light green color then intensi- L OTASSIUM Cu P . Plants grown with- fied to chlorosis in the middle and test difference between sample means was L F out K had tan-brown spots appear on upper leaves (Fig. 1G). Middle leaves Tissue nutrient concn (mg lower leaf margins, and K-deficient then began to develop a marginal, B

Mg), and sulfur deficiency ( plants had 38% less dry weight than the brown necrosis. Upper leaves then

L control plants. The K concentrations began to develop necrosis as the mid- DD indicates the of control plants and plants grown dle leaves became fully necrotic. NS under the K-deficient regime were BORON. Deficiency of B initially

B 6.07% and 0.47%, respectively. Over manifested in the young leaves. They L Mn), molybdenum deficiency ( time, these spots increased in size and were narrow and strap-like, and small, L crossed veins. Middle and lower leaves brown specks formed near where the

S then developed necrotic, mottled leaf blade and petiole met (Fig. 2A). L 0.001, respectively; spotting that ranged in coloration Dry weights between the two treat- £ Ca), magnesium deficiency ( P

L from dark black-brown to light tan- ments were not significantly different. green (Fig. 1D). The necrosis then Control tissue contained 56.0 mgÁkg–1

Mg covered the entire leaf. B, and tissue from plants grown un- 0.01, or L £ CALCIUM. Young leaves on der B-deficient conditions contained

P –1 Fe), manganese deficiency ( plants grown under Ca-deficient con- 13.0 mgÁkg B. As the disorder pro- L

0.05, ditions exhibited dark necrotic spots gressed, specks expanded into larger z £ on the leaf margin and downward curl dark brown or black spots. Spotting P K), calcium deficiency ( Ca

L (Fig. 1E). Plants grown without Ca also began to appear in the upper leaf L

test at had 37% less dry weight than those petioles. The young foliage on lower F plants that received a complete fertil- axillary shoots began to manifest the izer regime. Control plants had a Ca spotting. Lower axillary shoot buds Cu), iron deficiency (

L concentration of 1.66%, while plants died, and then the apical bud died. In K grown without Ca had a concentra- the advanced stage, lower leaves man- L tion of 0.40% Ca. Gray spots then ifested interveinal chlorosis that pro-

P), potassium deficiency ( appeared on the most recently ma- gressed to a brown coloration. Middle L

Tissue nutrient concn (%) tured leaf. The spots on both upper and upper leaves then manifested these and middle foliage then increased in symptoms. P

L size; some leaves became fully ne- Lower leaves exhibiting brown crotic. The apical bud became ne- spots along the margin was the initial crotic, and plant death followed. symptom of B toxicity (Fig. 2B). Dry MAGNESIUM. Plants grown un- weights of the control plants and der Mg-deficient conditions first plants grown under B-toxic condi- N * *** *** *** *** *** *** *** *** * *** NS NS

L exhibited gray-brown spots on the tions were not significantly different, N), phosphorus deficiency ( L B), boron toxicity (++B), copper deficiency (

2.38–5.61 0.30–1.24 2.90–5.86 1.00–2.50 0.25–0.86 0.73 25–75interveinal 25–75 7–35 60–300 regions 30–284 0.20–1.08 25–115 of middle leaves. but tissue B concentration was. The L A significant difference was not ob- control tissue B concentration was served between dry weights of the 46.8 mgÁkg–1, while the B-toxic tissue control plants and plants grown with- concentration was 242.1 mgÁkg–1.As out Mg. Differences in Mg tissue the disorder progressed, the spots

t concentration were significantly dif- spread along the margin. The mar- ferent with control plants containing ginal spotting moved into the middle 0.66% Mg compared with 0.08% Mg and upper leaves on the plant. u for the plants grown without Mg. As COPPER. Plants grown under Cu the disorder progressed, the size of deficiency exhibited a green-gray value Only one sample used in the number presented. *, **, or *** indicate statistically significant differences between sample means based on Treatments for nitrogen deficiency ( Asymptomatic treatment. Treatments for boron deficiency ( Insufficient sample was available to determine concentration. Mills and Jones, 1996. for scarlet sage. Element Table 3. ‘Evolution’ mealy-cup sage plant tissue nutrient concentrations as affected by deficient or toxic nutrient treatments and published optimu z y x w v u t Optimum range or value for scarlet sage Complete control 5.06 0.67 6.07 1.66 0.66 0.45 56.0 46.8 5.5 72.0 114.4 0.6 TreatmentP 2.57the 0.12 spots increased 0.47 0.40 (Fig. 1F), 0.08 and 0.24 the 13.0 242.1haze 1.1 coloration 55.5 7.6 that developed — on

506 • August 2012 22(4) middle and upper leaves (Fig. 2C). No and these asymptomatic plants were dry weight, leaf puckering and cup- significant difference was observed in sampled and analyzed for dry masses ping, and marginal and interveinal plant dry weights. Control Cu tissue and tissue concentrations to determine chlorosis that progressed to necrosis. concentration was 5.5 mgÁkg–1; tissue if nonvisual differences were evident. The gray discoloration we observed Cu concentration in plants grown There was no significant difference in was not mentioned. The lack of de- without Cu was 1.1 mgÁkg–1.Over dry weights for control plants and velopment of inflorescences in Zn- time, the haze discoloration encom- those grown under Mo-deficient con- deficient plants was also a unique passed the entire upper leaf surface. ditions. One control sample had a Mo symptom; while we observed smaller IRON. The development of a light concentration of 0.6 mgÁkg–1, while internodes, distortions of inflores- green coloration in the plants was Mo levels in all other plants including cences were typically associated with the first symptom of Fe deficiency other control replicate plants and Mo- B- and Ca-deficient conditions (Gibson (Fig. 2D). The coloration was not deficient plants were below the detect- et al., 2007). These unique symp- very apparent and would be difficult able limit. toms support a species-specific ap- to distinguish without a control pres- proach for visual diagnosis of nutrient ent. Dry weights of control plants and Discussion disorders because of the possibility of plants grown without Fe were not Symptoms of N, P, K, Ca, S, and misidentification. significantly different. The Fe con- Fe deficiencies and B toxicity mani- Necrosis was a common symp- centration of the control plants was fested earlier than other symptoms tom for several of the deficiencies (N, 72.0 mgÁkg–1, while plants grown (data not shown); therefore, these K, Ca, Mg, S, Mn, and Zn). Examin- under Fe-deficient conditions had disorders may be encountered more ing other symptoms would provide a concentration of 55.5 mgÁkg–1.As often by growers. The symptoms that more evidence for a diagnosis of the disorder progressed, some upper we observed for P, K, Ca, S, B, Fe, and a specific nutritional disorder. Early leaves developed a faint interveinal Mn deficiencies and B toxicity spotting observed with K, Ca, and Mg chlorosis. matched nutrient-disorder descrip- deficiencies led to necrosis. Both K and MANGANESE. Plants grown with- tions that were published in the liter- Ca deficiencies exhibited less chlorosis out Mn initially developed an overall ature (Barker and Pilbeam, 2007; than Mg deficiency. Less growth was light green coloration (Fig. 2E). Dry Gibson et al., 2007). For Mo de- observed with Ca-deficient plants than weights from the control treatment ficiency, symptoms may have not K- and Mg-deficient plants, and Ca- plants and plants grown under Mn- manifested because of low sensitivity deficient plants eventually collapsed. deficient conditions were not signifi- of the crop to the disorder, or the plug Necrosis appeared as mottled spotting cantly different. Control tissue had substrate provided adequate levels. with K-deficient plants instead of a Mn concentration of 114.4 mgÁkg–1, For N, Mg, Cu, and Zn, we interveinal necrosis observed with Mg and plants grown under Mn-deficient observed symptoms that were deficiency. Necrosis manifested in N, conditions contained 7.6 mgÁkg–1 Mn. unique. Leaf purpling occurred on S, Mn, and Zn deficiencies after other The light green coloration intensified N-deficient plants and not P-deficient symptoms appeared. Plants grown in to interveinal chlorosis in some leaves. plants in maize (Zea mays) and to- N-deficient conditions exhibited chlo- Brown spots then appeared on leaves mato (Solanum lycopersicum) (Bould rosis with purple coloration before throughout the plant either along the et al., 1984). This purpling of lower necrosis manifested on lower leaf tips. margin or in the interveinal areas; chlorotic leaves on N-deficient In S-deficient plants, chlorosis mani- symptoms were more pronounced plants was also reported for begonia fested before necrosis. Plants grown in on the middle leaves. The spotting (Begonia ·semperflorens-cultorum), Mn-deficient conditions exhibited sig- then progressed to leaf necrosis. celosia (Celosia argentea), marigold nificant growth before they first mani- ZINC. Upper leaves on plants (Tagetes patula), and pansy (Viola · fested chlorosis that then developed grown under Zn deficiency developed wittrockiana) (Gibson et al., 2007). into necrosis. For Zn deficiency, ne- a light green coloration (Fig. 2F). ForplantsgrowninMgdeficiency crotic spots developed on upper leaves Differences in Zn tissue concentra- conditions, we observed an in- with inflorescences that lacked full tions were significantly different at P = terveinal necrosis, while SePRO development. 0.06. The control tissue Zn concen- (2000) reported that interveinal In the macronutrient deficiency tration was 21.4 mgÁkg–1, while tissue chlorosis occurred for Mg defi- treatments, Na or Cl was used as the concentration in plants grown with- ciency. The symptoms observed for counterion. The highest level of Cl out Zn was 12.1 mgÁkg–1. As the dis- Cu-deficient plants—green-gray col- in solution was 531 mgÁkg–1 for the order progressed, the upper leaves oration on the inner portion of the N-deficiency treatment, which was began to develop brown, necrotic leaf blade—were the most unique within the sufficiency range of spots. Also, the inflorescence elonga- observed in this experiment because 20–1500 mgÁkg–1 reported by Jones tion and development were inhibited. they have not been described in (2000); further, Jones (2000) stated Axillary shoots lacked full develop- the literature. Gray coloration on that Cl is in excess after reaching levels ment as well. In the advanced stage, the tips of barley (Hordeum vulgare) of 5000 mgÁkg–1. For Na, we calcu- lower leaves exhibited chlorosis. and sunflower (Helianthus annuus) lated a sodium adsorption ratio (SAR) MOLYBDENUM. After 7 weeks of leaves was reported (Reuther and value of 0.67 for the solution; values <3 growth when the plants had reached Labanauskas, 1973). Gibson et al. are considered ideal, and no negative full bloom, plants grown under (2007) reports Cu-deficient symptoms impact is expected with <6 (Peterson, Mo deficiency exhibited no visual for scarlet sage. Symptoms included 1996). Therefore, it was unlikely that symptoms. The experiment was ended, Cu-deficient plants having reduced their elevated levels induced a Na

• August 2012 22(4) 507 RESEARCH REPORTS or Cl toxicity that confounded the Armitage, A., T. Dudek, S. Gortsema, and J.M. Dole. 2007. Nutrient deficiencies in results. B. Knox. 1994. Growing basics for salvia. bedding plants. Ball Publishing, Batavia, IL. When the control and defi- Professional Plant Growers Assn. News 25(5):17. Hoagland, R.J. and D.I. Arnon. 1950. cient treatment concentrations were The water-culture method for growing compared with Mills and Jones Barker, A.V. and D.J. Pilbeam. 2007. plants without soil. California Agr. Expt. (1996) optimum ranges and values Handbook of . CRC Press, Sta. Circ. 347 (revised ed.). (Table 3), for the control plants, eight Boca Raton, FL. Jones, J.B., Jr. 2000. : A concentration averages (N, P, Ca, Barnes, J. 2010. Characterization of nu- practical guide for the soilless grower. Mg, B, Fe, Mn, and Mo) fell within trient disorders in floriculture crops. St. Lucie Press, Boca Raton, FL. those reported values, three concen- North Carolina State Univ., Raleigh, MS tration averages (S, Cu, and Zn) were Thesis. Mills, H.A. and J.B. Jones, Jr. 1996. Plant below the values, and the K concen- analysis handbook II. Micromacro Pub- tration average was above the values. Bould, C., E.J. Hewitt, and P. Needham. lishing, Athens, GA. 1984. Diagnosis of mineral disorders in All of the deficient treatment values plants. Vol. 1. Chemical Publishing, Nelson, P. 2012. Greenhouse operation we reported except for N, which was New York. and management. 7th ed. Prentice Hall, in the lower end of their optimum Upper Saddle River, NJ. range, were lower than those reported Briskin, D.P. and A. Bloom. 2010. Min- eral nutrition, p. 107–130. In: L. Taiz and Peterson, F.H. 1996. Water testing and by Mills and Jones (1996). Our values interpretation, p. 31–49. In: D.W. Reed provide guidelines for interpreting E. Zeiger (eds.). Plant physiology. 5th ed. Sinauer Assoc., Sunderland, MA. (ed.). Water, media, and nutrition for the lower threshold tissue concentra- greenhouse crops. Ball Publishing, Batavia, tions, and the information reported Dole, J.M. and H.F. Wilkins. 2005. Flo- IL. riculture principles and species. 2nd ed. will be helpful to growers diagnosing Reuther, W. and C.K. Labanauskas. mealy-cup sage and other sage species Pearson-Prentice Hall, Upper Saddle River, NJ. 1973. Copper, p. 157–179. In: H.D. nutritional problems. Chapman (ed.). Diagnostic criteria for Dreistadt, S.H. 2001. Integrated pest plants and soils. Quality Printing Co., management for floriculture and nurseries. Abilene, TX. Univ. of California Publ. 3402. Literature cited SePRO. 2000. Salvia (sage): Cultural tips Armitage, A.M. 2001. Armitage’s manual Gibson, J.L., D.S. Pitchay, A.L. Williams- & guidelines. SePRO Corp. Carmel, IN. of annuals, biennials, and half-hardy pe- Rhodes, B.E. Whipker, P.V. Nelson, and rennials. Timber Press, Portland, OR.

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