Removal of nitrogen during needle senescence in Scots pine (Pinus sylvestris L.)
Abstract The concentrations of arginine, protein and total nitrogen (N) and the abundance of15N were measured in 3-and 4-year-old needles of Scots pine trees fertilized with either 0 (C), 36 (N1) or 73 (N2) kg N ha-1 year-1 annually for 22 years (average doses of N). Remaining green needles and need...
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| Format: |
E-Artikel |
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| Sprache: |
Englisch |
| Erschienen: |
1994 |
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| Umfang: |
7 |
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| Reproduktion: |
Springer Online Journal Archives 1860-2002 |
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| Übergeordnetes Werk: |
in: Oecologia - 1968, 99(1994) vom: März/Apr., Seite 290-296 |
| Übergeordnetes Werk: |
volume:99 ; year:1994 ; month:03/04 ; pages:290-296 ; extent:7 |
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NLEJ20590971X |
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| 520 | |a Abstract The concentrations of arginine, protein and total nitrogen (N) and the abundance of15N were measured in 3-and 4-year-old needles of Scots pine trees fertilized with either 0 (C), 36 (N1) or 73 (N2) kg N ha-1 year-1 annually for 22 years (average doses of N). Remaining green needles and needles that were shed were compared and removal of N from total, protein and arginine pools was calculated. Earlier investigations had shown that high arginine concentrations are found in needles of trees that have an excessive N supply (Näsholm and Ericsson 1990). This study aimed to elucidate the fate of the accumulated arginine during needle senescence. It was speculated that a low removal of arginine during senescence would implicate that the primary function of arginine is in N detoxification and not in N storage. Moreover, litter quality would be altered if needles are shed with high concentrations of arginine and this might affect the turnover of N in forest ecosystems. In remaining green needles, the concentration of total N increased with increasing N supply. Protein N concentrations were higher in fertilized trees, but did not differ between the two N treatments. Arginine N was low in C and N1 trees but high in N2 trees. Senescent needles from C and N1 trees had about equal total N concentrations while in N2 trees this concentration was significantly higher. Protein N in senescent needles did not differ between treatments. Arginine N, however, was less than 0.1 mg g−1 dw in C and N1 trees but was higher than 1.5 mg g−1 dw in N2 trees. Removal of N was highest in N1 trees followed by C trees while N2 trees removed least N from senescing needles. The high concentration of total N in senescent needles from N2 trees was to a great extent explained by a high arginine concentration. The δ15N value of remaining, green needles was higher (less negative) in N2 trees than in C and N1 trees. The same pattern was found for senescent needles. Comparisons of δ15N values between remaining, green and senescent needles within each treatment showed a significant increase in δ15N for all treatments during senescence possibly indicating losses of N as NH3 (g) from needles during senescence. It is concluded that arginine, accumulated in response to high N supply, is retranslocated only to a small extent during needle senescence. The ecological and physiological implications of this finding are discussed. | ||
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(DE-627)NLEJ20590971X DE-627 ger DE-627 rakwb eng Removal of nitrogen during needle senescence in Scots pine (Pinus sylvestris L.) 1994 7 nicht spezifiziert zzz rdacontent nicht spezifiziert z rdamedia nicht spezifiziert zu rdacarrier Abstract The concentrations of arginine, protein and total nitrogen (N) and the abundance of15N were measured in 3-and 4-year-old needles of Scots pine trees fertilized with either 0 (C), 36 (N1) or 73 (N2) kg N ha-1 year-1 annually for 22 years (average doses of N). Remaining green needles and needles that were shed were compared and removal of N from total, protein and arginine pools was calculated. Earlier investigations had shown that high arginine concentrations are found in needles of trees that have an excessive N supply (Näsholm and Ericsson 1990). This study aimed to elucidate the fate of the accumulated arginine during needle senescence. It was speculated that a low removal of arginine during senescence would implicate that the primary function of arginine is in N detoxification and not in N storage. Moreover, litter quality would be altered if needles are shed with high concentrations of arginine and this might affect the turnover of N in forest ecosystems. In remaining green needles, the concentration of total N increased with increasing N supply. Protein N concentrations were higher in fertilized trees, but did not differ between the two N treatments. Arginine N was low in C and N1 trees but high in N2 trees. Senescent needles from C and N1 trees had about equal total N concentrations while in N2 trees this concentration was significantly higher. Protein N in senescent needles did not differ between treatments. Arginine N, however, was less than 0.1 mg g−1 dw in C and N1 trees but was higher than 1.5 mg g−1 dw in N2 trees. Removal of N was highest in N1 trees followed by C trees while N2 trees removed least N from senescing needles. The high concentration of total N in senescent needles from N2 trees was to a great extent explained by a high arginine concentration. The δ15N value of remaining, green needles was higher (less negative) in N2 trees than in C and N1 trees. The same pattern was found for senescent needles. Comparisons of δ15N values between remaining, green and senescent needles within each treatment showed a significant increase in δ15N for all treatments during senescence possibly indicating losses of N as NH3 (g) from needles during senescence. It is concluded that arginine, accumulated in response to high N supply, is retranslocated only to a small extent during needle senescence. The ecological and physiological implications of this finding are discussed. Springer Online Journal Archives 1860-2002 Näsholm, Torgny oth in Oecologia 1968 99(1994) vom: März/Apr., Seite 290-296 (DE-627)NLEJ18898738X (DE-600)1462019-4 1432-1939 nnns volume:99 year:1994 month:03/04 pages:290-296 extent:7 http://dx.doi.org/10.1007/BF00627741 GBV_USEFLAG_U ZDB-1-SOJ GBV_NL_ARTICLE AR 99 1994 3/4 290-296 7 |
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(DE-627)NLEJ20590971X DE-627 ger DE-627 rakwb eng Removal of nitrogen during needle senescence in Scots pine (Pinus sylvestris L.) 1994 7 nicht spezifiziert zzz rdacontent nicht spezifiziert z rdamedia nicht spezifiziert zu rdacarrier Abstract The concentrations of arginine, protein and total nitrogen (N) and the abundance of15N were measured in 3-and 4-year-old needles of Scots pine trees fertilized with either 0 (C), 36 (N1) or 73 (N2) kg N ha-1 year-1 annually for 22 years (average doses of N). Remaining green needles and needles that were shed were compared and removal of N from total, protein and arginine pools was calculated. Earlier investigations had shown that high arginine concentrations are found in needles of trees that have an excessive N supply (Näsholm and Ericsson 1990). This study aimed to elucidate the fate of the accumulated arginine during needle senescence. It was speculated that a low removal of arginine during senescence would implicate that the primary function of arginine is in N detoxification and not in N storage. Moreover, litter quality would be altered if needles are shed with high concentrations of arginine and this might affect the turnover of N in forest ecosystems. In remaining green needles, the concentration of total N increased with increasing N supply. Protein N concentrations were higher in fertilized trees, but did not differ between the two N treatments. Arginine N was low in C and N1 trees but high in N2 trees. Senescent needles from C and N1 trees had about equal total N concentrations while in N2 trees this concentration was significantly higher. Protein N in senescent needles did not differ between treatments. Arginine N, however, was less than 0.1 mg g−1 dw in C and N1 trees but was higher than 1.5 mg g−1 dw in N2 trees. Removal of N was highest in N1 trees followed by C trees while N2 trees removed least N from senescing needles. The high concentration of total N in senescent needles from N2 trees was to a great extent explained by a high arginine concentration. The δ15N value of remaining, green needles was higher (less negative) in N2 trees than in C and N1 trees. The same pattern was found for senescent needles. Comparisons of δ15N values between remaining, green and senescent needles within each treatment showed a significant increase in δ15N for all treatments during senescence possibly indicating losses of N as NH3 (g) from needles during senescence. It is concluded that arginine, accumulated in response to high N supply, is retranslocated only to a small extent during needle senescence. The ecological and physiological implications of this finding are discussed. Springer Online Journal Archives 1860-2002 Näsholm, Torgny oth in Oecologia 1968 99(1994) vom: März/Apr., Seite 290-296 (DE-627)NLEJ18898738X (DE-600)1462019-4 1432-1939 nnns volume:99 year:1994 month:03/04 pages:290-296 extent:7 http://dx.doi.org/10.1007/BF00627741 GBV_USEFLAG_U ZDB-1-SOJ GBV_NL_ARTICLE AR 99 1994 3/4 290-296 7 |
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(DE-627)NLEJ20590971X DE-627 ger DE-627 rakwb eng Removal of nitrogen during needle senescence in Scots pine (Pinus sylvestris L.) 1994 7 nicht spezifiziert zzz rdacontent nicht spezifiziert z rdamedia nicht spezifiziert zu rdacarrier Abstract The concentrations of arginine, protein and total nitrogen (N) and the abundance of15N were measured in 3-and 4-year-old needles of Scots pine trees fertilized with either 0 (C), 36 (N1) or 73 (N2) kg N ha-1 year-1 annually for 22 years (average doses of N). Remaining green needles and needles that were shed were compared and removal of N from total, protein and arginine pools was calculated. Earlier investigations had shown that high arginine concentrations are found in needles of trees that have an excessive N supply (Näsholm and Ericsson 1990). This study aimed to elucidate the fate of the accumulated arginine during needle senescence. It was speculated that a low removal of arginine during senescence would implicate that the primary function of arginine is in N detoxification and not in N storage. Moreover, litter quality would be altered if needles are shed with high concentrations of arginine and this might affect the turnover of N in forest ecosystems. In remaining green needles, the concentration of total N increased with increasing N supply. Protein N concentrations were higher in fertilized trees, but did not differ between the two N treatments. Arginine N was low in C and N1 trees but high in N2 trees. Senescent needles from C and N1 trees had about equal total N concentrations while in N2 trees this concentration was significantly higher. Protein N in senescent needles did not differ between treatments. Arginine N, however, was less than 0.1 mg g−1 dw in C and N1 trees but was higher than 1.5 mg g−1 dw in N2 trees. Removal of N was highest in N1 trees followed by C trees while N2 trees removed least N from senescing needles. The high concentration of total N in senescent needles from N2 trees was to a great extent explained by a high arginine concentration. The δ15N value of remaining, green needles was higher (less negative) in N2 trees than in C and N1 trees. The same pattern was found for senescent needles. Comparisons of δ15N values between remaining, green and senescent needles within each treatment showed a significant increase in δ15N for all treatments during senescence possibly indicating losses of N as NH3 (g) from needles during senescence. It is concluded that arginine, accumulated in response to high N supply, is retranslocated only to a small extent during needle senescence. The ecological and physiological implications of this finding are discussed. Springer Online Journal Archives 1860-2002 Näsholm, Torgny oth in Oecologia 1968 99(1994) vom: März/Apr., Seite 290-296 (DE-627)NLEJ18898738X (DE-600)1462019-4 1432-1939 nnns volume:99 year:1994 month:03/04 pages:290-296 extent:7 http://dx.doi.org/10.1007/BF00627741 GBV_USEFLAG_U ZDB-1-SOJ GBV_NL_ARTICLE AR 99 1994 3/4 290-296 7 |
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(DE-627)NLEJ20590971X DE-627 ger DE-627 rakwb eng Removal of nitrogen during needle senescence in Scots pine (Pinus sylvestris L.) 1994 7 nicht spezifiziert zzz rdacontent nicht spezifiziert z rdamedia nicht spezifiziert zu rdacarrier Abstract The concentrations of arginine, protein and total nitrogen (N) and the abundance of15N were measured in 3-and 4-year-old needles of Scots pine trees fertilized with either 0 (C), 36 (N1) or 73 (N2) kg N ha-1 year-1 annually for 22 years (average doses of N). Remaining green needles and needles that were shed were compared and removal of N from total, protein and arginine pools was calculated. Earlier investigations had shown that high arginine concentrations are found in needles of trees that have an excessive N supply (Näsholm and Ericsson 1990). This study aimed to elucidate the fate of the accumulated arginine during needle senescence. It was speculated that a low removal of arginine during senescence would implicate that the primary function of arginine is in N detoxification and not in N storage. Moreover, litter quality would be altered if needles are shed with high concentrations of arginine and this might affect the turnover of N in forest ecosystems. In remaining green needles, the concentration of total N increased with increasing N supply. Protein N concentrations were higher in fertilized trees, but did not differ between the two N treatments. Arginine N was low in C and N1 trees but high in N2 trees. Senescent needles from C and N1 trees had about equal total N concentrations while in N2 trees this concentration was significantly higher. Protein N in senescent needles did not differ between treatments. Arginine N, however, was less than 0.1 mg g−1 dw in C and N1 trees but was higher than 1.5 mg g−1 dw in N2 trees. Removal of N was highest in N1 trees followed by C trees while N2 trees removed least N from senescing needles. The high concentration of total N in senescent needles from N2 trees was to a great extent explained by a high arginine concentration. The δ15N value of remaining, green needles was higher (less negative) in N2 trees than in C and N1 trees. The same pattern was found for senescent needles. Comparisons of δ15N values between remaining, green and senescent needles within each treatment showed a significant increase in δ15N for all treatments during senescence possibly indicating losses of N as NH3 (g) from needles during senescence. It is concluded that arginine, accumulated in response to high N supply, is retranslocated only to a small extent during needle senescence. The ecological and physiological implications of this finding are discussed. Springer Online Journal Archives 1860-2002 Näsholm, Torgny oth in Oecologia 1968 99(1994) vom: März/Apr., Seite 290-296 (DE-627)NLEJ18898738X (DE-600)1462019-4 1432-1939 nnns volume:99 year:1994 month:03/04 pages:290-296 extent:7 http://dx.doi.org/10.1007/BF00627741 GBV_USEFLAG_U ZDB-1-SOJ GBV_NL_ARTICLE AR 99 1994 3/4 290-296 7 |
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(DE-627)NLEJ20590971X DE-627 ger DE-627 rakwb eng Removal of nitrogen during needle senescence in Scots pine (Pinus sylvestris L.) 1994 7 nicht spezifiziert zzz rdacontent nicht spezifiziert z rdamedia nicht spezifiziert zu rdacarrier Abstract The concentrations of arginine, protein and total nitrogen (N) and the abundance of15N were measured in 3-and 4-year-old needles of Scots pine trees fertilized with either 0 (C), 36 (N1) or 73 (N2) kg N ha-1 year-1 annually for 22 years (average doses of N). Remaining green needles and needles that were shed were compared and removal of N from total, protein and arginine pools was calculated. Earlier investigations had shown that high arginine concentrations are found in needles of trees that have an excessive N supply (Näsholm and Ericsson 1990). This study aimed to elucidate the fate of the accumulated arginine during needle senescence. It was speculated that a low removal of arginine during senescence would implicate that the primary function of arginine is in N detoxification and not in N storage. Moreover, litter quality would be altered if needles are shed with high concentrations of arginine and this might affect the turnover of N in forest ecosystems. In remaining green needles, the concentration of total N increased with increasing N supply. Protein N concentrations were higher in fertilized trees, but did not differ between the two N treatments. Arginine N was low in C and N1 trees but high in N2 trees. Senescent needles from C and N1 trees had about equal total N concentrations while in N2 trees this concentration was significantly higher. Protein N in senescent needles did not differ between treatments. Arginine N, however, was less than 0.1 mg g−1 dw in C and N1 trees but was higher than 1.5 mg g−1 dw in N2 trees. Removal of N was highest in N1 trees followed by C trees while N2 trees removed least N from senescing needles. The high concentration of total N in senescent needles from N2 trees was to a great extent explained by a high arginine concentration. The δ15N value of remaining, green needles was higher (less negative) in N2 trees than in C and N1 trees. The same pattern was found for senescent needles. Comparisons of δ15N values between remaining, green and senescent needles within each treatment showed a significant increase in δ15N for all treatments during senescence possibly indicating losses of N as NH3 (g) from needles during senescence. It is concluded that arginine, accumulated in response to high N supply, is retranslocated only to a small extent during needle senescence. The ecological and physiological implications of this finding are discussed. Springer Online Journal Archives 1860-2002 Näsholm, Torgny oth in Oecologia 1968 99(1994) vom: März/Apr., Seite 290-296 (DE-627)NLEJ18898738X (DE-600)1462019-4 1432-1939 nnns volume:99 year:1994 month:03/04 pages:290-296 extent:7 http://dx.doi.org/10.1007/BF00627741 GBV_USEFLAG_U ZDB-1-SOJ GBV_NL_ARTICLE AR 99 1994 3/4 290-296 7 |
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Remaining green needles and needles that were shed were compared and removal of N from total, protein and arginine pools was calculated. Earlier investigations had shown that high arginine concentrations are found in needles of trees that have an excessive N supply (Näsholm and Ericsson 1990). This study aimed to elucidate the fate of the accumulated arginine during needle senescence. It was speculated that a low removal of arginine during senescence would implicate that the primary function of arginine is in N detoxification and not in N storage. Moreover, litter quality would be altered if needles are shed with high concentrations of arginine and this might affect the turnover of N in forest ecosystems. In remaining green needles, the concentration of total N increased with increasing N supply. Protein N concentrations were higher in fertilized trees, but did not differ between the two N treatments. Arginine N was low in C and N1 trees but high in N2 trees. Senescent needles from C and N1 trees had about equal total N concentrations while in N2 trees this concentration was significantly higher. Protein N in senescent needles did not differ between treatments. Arginine N, however, was less than 0.1 mg g−1 dw in C and N1 trees but was higher than 1.5 mg g−1 dw in N2 trees. Removal of N was highest in N1 trees followed by C trees while N2 trees removed least N from senescing needles. The high concentration of total N in senescent needles from N2 trees was to a great extent explained by a high arginine concentration. The δ15N value of remaining, green needles was higher (less negative) in N2 trees than in C and N1 trees. The same pattern was found for senescent needles. Comparisons of δ15N values between remaining, green and senescent needles within each treatment showed a significant increase in δ15N for all treatments during senescence possibly indicating losses of N as NH3 (g) from needles during senescence. It is concluded that arginine, accumulated in response to high N supply, is retranslocated only to a small extent during needle senescence. 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removal of nitrogen during needle senescence in scots pine (pinus sylvestris l.) |
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Removal of nitrogen during needle senescence in Scots pine (Pinus sylvestris L.) |
| abstract |
Abstract The concentrations of arginine, protein and total nitrogen (N) and the abundance of15N were measured in 3-and 4-year-old needles of Scots pine trees fertilized with either 0 (C), 36 (N1) or 73 (N2) kg N ha-1 year-1 annually for 22 years (average doses of N). Remaining green needles and needles that were shed were compared and removal of N from total, protein and arginine pools was calculated. Earlier investigations had shown that high arginine concentrations are found in needles of trees that have an excessive N supply (Näsholm and Ericsson 1990). This study aimed to elucidate the fate of the accumulated arginine during needle senescence. It was speculated that a low removal of arginine during senescence would implicate that the primary function of arginine is in N detoxification and not in N storage. Moreover, litter quality would be altered if needles are shed with high concentrations of arginine and this might affect the turnover of N in forest ecosystems. In remaining green needles, the concentration of total N increased with increasing N supply. Protein N concentrations were higher in fertilized trees, but did not differ between the two N treatments. Arginine N was low in C and N1 trees but high in N2 trees. Senescent needles from C and N1 trees had about equal total N concentrations while in N2 trees this concentration was significantly higher. Protein N in senescent needles did not differ between treatments. Arginine N, however, was less than 0.1 mg g−1 dw in C and N1 trees but was higher than 1.5 mg g−1 dw in N2 trees. Removal of N was highest in N1 trees followed by C trees while N2 trees removed least N from senescing needles. The high concentration of total N in senescent needles from N2 trees was to a great extent explained by a high arginine concentration. The δ15N value of remaining, green needles was higher (less negative) in N2 trees than in C and N1 trees. The same pattern was found for senescent needles. Comparisons of δ15N values between remaining, green and senescent needles within each treatment showed a significant increase in δ15N for all treatments during senescence possibly indicating losses of N as NH3 (g) from needles during senescence. It is concluded that arginine, accumulated in response to high N supply, is retranslocated only to a small extent during needle senescence. The ecological and physiological implications of this finding are discussed. |
| abstractGer |
Abstract The concentrations of arginine, protein and total nitrogen (N) and the abundance of15N were measured in 3-and 4-year-old needles of Scots pine trees fertilized with either 0 (C), 36 (N1) or 73 (N2) kg N ha-1 year-1 annually for 22 years (average doses of N). Remaining green needles and needles that were shed were compared and removal of N from total, protein and arginine pools was calculated. Earlier investigations had shown that high arginine concentrations are found in needles of trees that have an excessive N supply (Näsholm and Ericsson 1990). This study aimed to elucidate the fate of the accumulated arginine during needle senescence. It was speculated that a low removal of arginine during senescence would implicate that the primary function of arginine is in N detoxification and not in N storage. Moreover, litter quality would be altered if needles are shed with high concentrations of arginine and this might affect the turnover of N in forest ecosystems. In remaining green needles, the concentration of total N increased with increasing N supply. Protein N concentrations were higher in fertilized trees, but did not differ between the two N treatments. Arginine N was low in C and N1 trees but high in N2 trees. Senescent needles from C and N1 trees had about equal total N concentrations while in N2 trees this concentration was significantly higher. Protein N in senescent needles did not differ between treatments. Arginine N, however, was less than 0.1 mg g−1 dw in C and N1 trees but was higher than 1.5 mg g−1 dw in N2 trees. Removal of N was highest in N1 trees followed by C trees while N2 trees removed least N from senescing needles. The high concentration of total N in senescent needles from N2 trees was to a great extent explained by a high arginine concentration. The δ15N value of remaining, green needles was higher (less negative) in N2 trees than in C and N1 trees. The same pattern was found for senescent needles. Comparisons of δ15N values between remaining, green and senescent needles within each treatment showed a significant increase in δ15N for all treatments during senescence possibly indicating losses of N as NH3 (g) from needles during senescence. It is concluded that arginine, accumulated in response to high N supply, is retranslocated only to a small extent during needle senescence. The ecological and physiological implications of this finding are discussed. |
| abstract_unstemmed |
Abstract The concentrations of arginine, protein and total nitrogen (N) and the abundance of15N were measured in 3-and 4-year-old needles of Scots pine trees fertilized with either 0 (C), 36 (N1) or 73 (N2) kg N ha-1 year-1 annually for 22 years (average doses of N). Remaining green needles and needles that were shed were compared and removal of N from total, protein and arginine pools was calculated. Earlier investigations had shown that high arginine concentrations are found in needles of trees that have an excessive N supply (Näsholm and Ericsson 1990). This study aimed to elucidate the fate of the accumulated arginine during needle senescence. It was speculated that a low removal of arginine during senescence would implicate that the primary function of arginine is in N detoxification and not in N storage. Moreover, litter quality would be altered if needles are shed with high concentrations of arginine and this might affect the turnover of N in forest ecosystems. In remaining green needles, the concentration of total N increased with increasing N supply. Protein N concentrations were higher in fertilized trees, but did not differ between the two N treatments. Arginine N was low in C and N1 trees but high in N2 trees. Senescent needles from C and N1 trees had about equal total N concentrations while in N2 trees this concentration was significantly higher. Protein N in senescent needles did not differ between treatments. Arginine N, however, was less than 0.1 mg g−1 dw in C and N1 trees but was higher than 1.5 mg g−1 dw in N2 trees. Removal of N was highest in N1 trees followed by C trees while N2 trees removed least N from senescing needles. The high concentration of total N in senescent needles from N2 trees was to a great extent explained by a high arginine concentration. The δ15N value of remaining, green needles was higher (less negative) in N2 trees than in C and N1 trees. The same pattern was found for senescent needles. Comparisons of δ15N values between remaining, green and senescent needles within each treatment showed a significant increase in δ15N for all treatments during senescence possibly indicating losses of N as NH3 (g) from needles during senescence. It is concluded that arginine, accumulated in response to high N supply, is retranslocated only to a small extent during needle senescence. The ecological and physiological implications of this finding are discussed. |
| collection_details |
GBV_USEFLAG_U ZDB-1-SOJ GBV_NL_ARTICLE |
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Removal of nitrogen during needle senescence in Scots pine (Pinus sylvestris L.) |
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http://dx.doi.org/10.1007/BF00627741 |
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Näsholm, Torgny |
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2024-07-06T05:23:32.078Z |
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Remaining green needles and needles that were shed were compared and removal of N from total, protein and arginine pools was calculated. Earlier investigations had shown that high arginine concentrations are found in needles of trees that have an excessive N supply (Näsholm and Ericsson 1990). This study aimed to elucidate the fate of the accumulated arginine during needle senescence. It was speculated that a low removal of arginine during senescence would implicate that the primary function of arginine is in N detoxification and not in N storage. Moreover, litter quality would be altered if needles are shed with high concentrations of arginine and this might affect the turnover of N in forest ecosystems. In remaining green needles, the concentration of total N increased with increasing N supply. Protein N concentrations were higher in fertilized trees, but did not differ between the two N treatments. Arginine N was low in C and N1 trees but high in N2 trees. 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